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COVID-19 has wreaked havoc globally with particular concerns for sub-Saharan Africa (SSA), where models suggest that the majority of the population will become infected. Conventional wisdom suggests that the continent will bear a higher burden of COVID-19 for the same reasons it suffers from other infectious diseases: ecology, socio-economic conditions, lack of water and sanitation infrastructure, and weak health systems. However, so far SSA has reported lower incidence and fatalities compared to the predictions of standard models and the experience of other regions of the world. There are three leading explanations, each with different implications for the final epidemic burden: (1) low case detection, (2) differences in epidemiology (e.g. lowR(0)), and (3) policy interventions. The low number of cases have led some SSA governments to relaxing these policy interventions. Will this result in a resurgence of cases? To understand how to interpret the lower-than-expected COVID-19 case data in Madagascar, we use a simple age-structured model to explore each of these explanations and predict the epidemic impact associated with them. We show that the incidence of COVID-19 cases as of July 2020 can be explained by any combination of the late introduction of first imported cases, early implementation of non-pharmaceutical interventions (NPIs), and low case detection rates. We then re-evaluate these findings in the context of the COVID-19 epidemic in Madagascar through August 2020. This analysis reinforces that Madagascar, along with other countries in SSA, remains at risk of a growing health crisis. If NPIs remain enforced, up to 50,000 lives may be saved. Even with NPIs, without vaccines and new therapies, COVID-19 could infect up to 30% of the population, making it the largest public health threat in Madagascar for the coming year, hence the importance of clinical trials and continually improving access to healthcare.

Diagnosis of workers and queens of P. wasmannii-group species It should be noted that all diagnostic characters listed below are not unique to Malagasy P. wasmannii-group species but are shared with some Bothroponera (sensu Wheeler 1922) species from the Afrotropics. 1. Mandible subtriangular and robust, with masticatory margins bearing 7-9 teeth and denticles; groove running along the lateral portion from the base to the apex, but without a basilateral pit. 2. Palp formula 4,4. 3. Frontal lobes noticeably widely rounded anteriorly, the outer margins posteriorly narrow and compressed laterally at about the level of the eyes. 4. Clypeus usually with a truncate anteromedian margin, the shape of which is straight or slightly excised medially, but in P. tavaratra projects into a triangular lobe. The median portion posteriorly pinched into a slight triangle between the frontal lobes and extending backwards as a line at about the midlength of the head. 5. Antenna with 12 segments, the funiculus gradually increasing in width towards the apex and without a distinct club. 6. Compound eyes present and generally quite large but their maximum diameter less than the maximum width of the scape; posterior margin of the eye placed in the front of the midline of the head when viewed in profile. 7. Tibia of the hind legs with a pair of apical spurs, the posterior pectinate and the anterior smaller and simple. 8. Pretarsal claws generally simple, preapical tooth may be present at the base in larger species (P. comorensis, P. tavaratra). 9. Metanotal groove obsolete or lacking in the worker caste. 10. Mesopleural sulcus either clearly visible in P. masoala, P. wasmannii, and in some P. cambouei or indistinct in P. perroti, P. vazimba, and in some P. planicornis. 11. Propodeum generally broad in dorsal view, the rear margin rounded, not bidentate or bispinose. 12. Propodeal spiracle slit-like. 13. Metapleural gland opening just above the posteroventral angle of mesosoma. 14. Basal fourth of all femora strongly laterally flattened (Fig. 52). 15. Petiole nodiform, usually thick, tapered towards the apex in P. masoala. 16. Subpetiolar process hook-like anteriorly and with nearly flattened or very slightly convex surface; sometimes with transverse ridges in the posterior portion in P. comorensis, P. tavaratra, and in some P. cambouei. 17. Helcium almost on the anteroventral angle of the first gastral segment (abdominal segment IV). 18. Strong constriction between the first and second gastral segments. 19. Stridulitrum present. 20. Sting well developed. 21. Queen caste-specific characteristics: broader head, presence of ocelli, eyes larger than the greatest width of antennal scape; mesopleuron divided into an episternum and katepisternum by a transverse sulcus, the mesosomal flight sclerites fully developed; petiole node more flattened anteroposteriorly relative to that of worker; somewhat larger body size with usually larger gaster and denser and more elongate pubescence. In the Malagasy P. wasmannii-group species, the worker caste can generally be identified by the following character combination: mandibles without basi-lateral pit; obsolete metanotal groove; propodeal spiracle opening slit-shaped; hind leg with two tibial spurs, of which one is large and pectinate and the other smaller and simple; roughly basal fourth of all femora strongly laterally flattened; propodeal rear margin simple; petiolar node thick whithout spines on rear margin; and first and second gastral segments seperated by a strong constriction. In addition, the mesosoma and petiole node usually strongly sculptured, frontal lobes broadly rounded, and propodeal dorsum wide. Synopsis of the taxonomic history of the P. wasmannii species-group The four previously described species in the P. wasmannii-group were all described between 1887 and 1892 in the genus Bothroponera. We summarize the taxonomic history of Bothroponera below. Bothroponera Mayr, 1862:713, 717 [as subgenus of Ponera by Emery, 1895b: 767 and Forel, 1900: 322; as subgenus of Pachycondyla by Emery, 1901: 42; Wheeler, 1910: 135, 1911: 160, 1917: 489; Emery, 1911: 74, 75, 76; Arnold, 1915: 55; Forel, 1917: 237; Donisthorpe, 1943: 628. As genus by Forel, 1891: 127; Dalla Torre 1893: 35; Bingham, 1903: 95; Ashmead, 1905: 382; Wheeler 1918: 299, 1922: 1008; Taylor and Brown, 1985: 21; Tiwari, 1999: 11, 27, 28; Schmidt, 2009: 119 (PhD thesis). As junior synonym of Pachycondyla by Brown, 1973: 179 [provisional]; Snelling 1981: 389; Hölldobler and Wilson, 1990: 11; Bolton, 1994: 164; Bolton, 2003: 166]. Type species: Ponera pumicosa Roger, 1860: 290; by subsequent designation of Emery, 1901: 42. Synoptic checklist of Malagasy P. wasmannii-group species cambouei Forel 1891 comorensis André 1887 masoala Rakotonirina and Fisher, sp. n. perroti Forel 1891 = perroti admista Forel 1892syn. n. planicornis Rakotonirina and Fisher, sp. n. tavaratra Rakotonirina and Fisher, sp. n. vazimba Rakotonirina and Fisher, sp. n. wasmannii Forel 1887 Key to species of the Malagasy P. wasmannii-group worker: The following key also identifies ergatoid and winged queens to species as long as the form of the mesosoma and the presence of the mesopleural sulcus are not applied. 1. Larger species (HW: 2.49-3.13 mm); with head in full-face view, antennal scape surpassing the posterior margin of the head (SI: 94-104) (Figs 1, 3); dorsum of head and mesosoma finely striate and interspersed with scattered punctures; dorsum of the body covered with erect stout hairs (Figs 30, 38).............................................................2 Smaller species (HW: 1.19-2.58 mm); with head in full-face view, antennal scape not surpassing the posterior margin of the head (SI: 70-87) (Fig. 2); dorsum of head and mesosoma variably sculptured but not finely striate, dorsum of the body with erect slender hairs and pubescence (e.g., Figs 35, 43, 45)......................................................3 2. With head in full-face view, anterior margin of clypeus broadly rounded or medially triangular; eye breaking the outline of the side of the head (Fig. 3); hairs on the dorsum of the mesosoma and petiole node distinctly erect relative to the body surface (Fig. 47).......................................................................................tavaratra With head in full-face view, anterior margin of clypeus truncate; eye not breaking the outline of the side of the head (Fig. 4); hairs on the dorsum of the body decumbent and much more inclined to the body surface (Fig. 39)..............comorensis 3. Basal half of the antennal scape dorsoventrally flattened, the leading edge very thin (Fig. 5); basitarsus of the hind leg laterally compressed, basal half of its inner surface concave...................................................planicornis Basal half of the antennal scape subcylindrical, not dorsoventrally flattened; the leading edge rounded (Fig. 6); basitarsus of the hind leg generally rounded, without concavity............................................................4 4. With petiole in dorsal view, the node flattened, about twice as broad as long (Fig.7); higher than long in profile; erect hairs on dorsum of propodeum most frequently absent (Fig. 41), if present then reduced in number...................... masoala With petiole in dorsal view, the node thick, approximately as long as broad (Fig. 8), about as high as long in profile; erect hairs on dorsum of propodeum numerous (e.g., Figs 18, 43, 51) or rarely absent (Fig. 31).................................5 5. Anterior half of the fourth abdominal tergite (gastral tergite 2) covered with numerous and very dense small punctures between larger ones (Fig. 9) which become moderate on mesosoma and petiole node; integument usually matte; mostly found in dry forest habitats........................................................................................6 Anterior half of the fourth abdominal tergite (gastral tergite 2) mostly smooth and shiny between large punctures, sometimes with randomly scattered small punctures (Fig. 10); mesosoma and petiole node sparsely punctulate between coarse punctae; integument mostly shiny; humid forest species.............................................................. 7 6. Larger species (HW: 1.72-2.21 mm); mesopleural sulcus usually distinct and complete (Fig. 11); dorsum of mesosoma and petiole node covered with more robust and longer standing hairs (Fig. 51); antennal segments 6, 7, and 8 nearly as wide as long; with petiole in dorsal view, the posterior margin straight or with weak median notch; outer surface of the hind tibia with erect hairs................................................................................... wasmannii Smaller species (HW: 1.38-1.5 mm); mesopleural sulcus indistinct or incomplete (Fig. 12); dorsum of mesosoma and petiole node with short and thin erect hairs (Fig. 49); antennal segments 6, 7, and 8 nearly twice as wide as long (Fig. 48); with petiole in dorsal view, the posterior margin broadly concave; outer surface of the hind tibia usually without erect hairs..... vazimba sulcus of P. wasmannii complete; Fig. 12: mesopleural 7. With mesosoma in profile, the dorsal outline forming a continuous convexity with a nearly rounded junction between the propodeal dorsum and the declivitous margin (Fig. 13), which is short and strongly inclined posteriorly; in full-face view, head heart-shaped, with the posterior margin strongly concave, and anterior clypeal margin strongly excised medially (Fig. 42); larger species (HW: 2.34-2.58 mm).................................................................. perroti With mesosoma in profile, the dorsal outline not forming a continuous convexity, propodeal dorsum meeting the declivity at a distinct angle (Fig. 14); propodeal declivity nearly vertical, but not distinctly inclined posteriorly; in full-face view head not heart-shaped, the posterior margin usually straight (Figs 30, 34, 36) or slightly concave (Figs 26, 28, 32); smaller species (HW

Pachycondyla zoro Rakotonirina and Fisher, sp. nov. (Figures 21, 22, 67, 68,69, 81) Holotype worker: Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38�� NE Andapa, 8.2 km 333�� NNW Manantenina, -14.4367, 49.775, 450 m, sifted litter (leaf mold, rotten wood), rainforest, 12-15 Nov 2003 (coll. B.L. Fisher et al.), collection code BLF08722, specimen code CASENT0045620 (CASC). Paratypes: series of seven workers and one ergatoid queen with same data as holotype but specimens coded as CASENT0045613, CASENT0045615, CASENT0045616, CASENT0045618, CASENT0045619, CASENT0045623, CASENT0045627, CASENT0045614 (BMNH, CASC, MHNG, MCZC, PBZT). WORKER. Diagnosis: Anterior margin of clypeus medially convex or with median obtuse angle; with head in full-face view, lateral margins before level of anterior margin of eye strongly converging toward the base of mandibles; anterior surface of third abdominal segment straight, without shallow cavity. Posterolateral margins of propodeum and petiole node without succession of sharp teeth or denticles, posterior margin of propodeum bordered with subopaque lamella; in dorsal view, petiole node nearly as long as broad; in lateral view, subpetiolar process with anterior hook-like and posterior subtriangular processes separated by a concavity. Measurements (8 specimens): HW: 1.22-1.32, HL: 1.33-1.46, CI: 90-94, SL: 1.11-1.17, SI: 87-92, PW: 0.95-1.02, WL: 1.85-2.05, NH: 0.77-0.83, NL: 0.53-0.61, NW: 0.67-0.73, DNI: 115-136, LNI: 135-154. Description: In full-face view, head subrectangular; longer than broad, with almost straight lateral margins posterior to the level of eye. Anterior to the level of eye, lateral margin curves strongly toward the base of mandibles; posterior margin weakly concave. Frontal lobe narrow, with broadly rounded outer margin. Head dorsally finely reticulate-punctate, with smooth and shiny surface between punctures; side smooth and shiny apart from effaced reticulation and shallow scattered punctures. Eyes medium, their diameter about the same as greatest width of scape. Antennal scape barely extending beyond posterior cephalic margin. Median lobe of clypeus weakly protruding forwards, anterior margin medially convex and sometimes with feeble median notch. Mandible smooth between sparse piligerous punctulae, masticatory margin with 9-12 teeth or denticles. In dorsal view, metanotal groove obsolete; promesonotum rugulose-punctate with smooth and shiny surface towards midline of mesosoma; propodeal dorsum sparsely punctate. In lateral view, mesopleural suture indistinct, median portion of mesopleuron and metapleuron shining and nearly smooth apart from widely spaced punctures or foveolae, with rugulose sculpture near lateroventral angle; sides of propodeum covered with dense, elongate punctures; posterolateral margin of propodeum bordered with subopaque lamellae. Petiole node in lateral view higher than long, with rugulose surface; anterodorsal angle extending somewhat over anterior face; posterior margin convex; subpetiole with both anterior hook-like and posterior subquadrate processes separated by a concavity. In dorsal view, node as long as broad, with nearly smooth dorsum apart from scattered rugulae and shallow punctures. Shallow cavity absent from anterior surface of first gastral segment. First and second gastral tergites with spaced punctures from which erect hairs arise. Pilosity consists of golden short erect hairs and abundant pubescence on head and promesonotum; rest of body dorsum with long erect hairs and very little pubescence. Head, mesosoma and petiolar node black; appendages and gaster brown ferruginous. ERGATOID QUEEN. Measurements (1 specimen): HW: 1.25, HL: 1.28, CI: 98, SL: 1.08, SI: 86, PW: 0.99, WL: 1.91, NH: 0.81, NL: 0.51, NW: 0.79, DNI: 155, LNI: 159. This specimen is very similar to workers due to the absence of wings and thoracic sclerite development. It differs by the presence of one ocellus, the broader head and much more slender and numerous erect hairs on the dorsum of the body. In addition, the petiolar node looks shorter when viewed in profile and much broader than those of workers when in dorsal view. Distribution and biology: Pachycondyla zoro occupies the lowland rainforests of Marojejy and Ambalagoavy Nord (Ikongo) located in the northeast and southeast of Madagascar respectively (Fig. 81). It has been collected via leaf litter extraction in Marojejy, and from malaise traps in Ambalagoavy. Additional material examined: Province Antsiranana: PN Marojejy, Manantenina River, 28.0 km 38�� NE Andapa, 8.2 km 333�� NNW Manantenina, Antsiranana, -14.4367, 49.775, 450 m, rainforest (B.L. Fisher et al.) (CASC); Province Fianarantsoa: F d'Ambalagoavy Nord, Ikongo, Ambatombe, -21.8275, 47.3389, 625 m (R. Harin'Hala & M.E. Irwin) (CASC). Aknowledgments We greatly appreciate the help of B. Merz, S. Cover, C. Villemant, and P.S. Ward for providing type material and additional specimens from their collections. We are grateful to the arthropod team at the Madagascar Biodiversity Center for carrying out the ant surveys, laboratory processing, and specimen sorting. The Malagasy Pachycondyla database could not have been refined and made easily available without the gracious management of M. Esposito. The manuscript was greatly improved by comments provided by the reviewers. The present contribution was part of the MSc research of JCR supported by The Lakeside Foundation Funds and McBean Family Fund of the California Academy of Sciences. Additional funding also came from National Science Foundation grants DEB- 0072713, DEB-0344731, and DEB-0842395, awarded to BLF., Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 480-484

Pachycondyla perroti (Forel) (Figures 13, 42-43, 60) Bothroponera perroti Forel, 1891: 131 pl. 4, fig. 6. Holotype worker, Madagascar, Toamasina, Fenerive (E. Perrot), AntWeb specimen code: CASENT0101401 (MNHN) [examined]. [Further description by Dalla Torre 1893: 36. Combination in Ponera (Bothroponera):Emery, 1899: 267; in Pachycondyla (Bothroponera):Emery, 1901: 45; Combination in Bothroponera: Wheeler, 1922: 1007; in Pachycondyla:Bolton, 1995: 308]. Bothroponera perroti admista Forel, 1892: 251. Lectotype worker, present designation Madagascar, Anosibe Bezanozano Province Moramanga (Sikora), AntWeb specimen code: CASENT0101035 (MHNG) [examined]. Syn. n. Paralectotype worker with the same data but specimen coded as CASENT0101034 (MHNG) [examined]. [Raised to species: Dalla Torre, 1893: 35; returned to subspecies:Wasmann, 1897: 250, Forel, 1897: 188, Emery, 1901: 45. Combination in Ponera (Bothroponera): Emery, 1899: 267; in Pachycondyla (Bothroponera): Emery, 1901: 45, 1911: 78; Combination in Bothroponera: Wheeler, 1922: 1008; in Pachycondyla: Bolton, 1995: 302]. Worker diagnosis: With head in full-face view, antennal scape not surpassing posterior cephalic margin; dorsum of head and mesosoma not finely striate, dorsum of the body with erect slender hairs and pubescence; basal half of antennal scape rounded; dorsal outline of mesosoma forming a continuous convexity with a nearly rounded junction between propodeal dorsum and declivitous surface; anterior half of fourth abdominal tergite (gastral tergite 2) mostly smooth and shiny between large punctures. Worker measurements (n=26): HL: 2.42-2.71, HW: 2.27-2.58, CI: 91-98, SL: 1.69-2.01, SI: 69-81, PW: 1.58-1.81, WL: 3.10-3.55, NL: 0.87-1.04, NW: 1.02-1.23, NH: 1.18-1.45, DNI: 108-130, LNI: 125-144. Description Worker. Head as long as broad, but narrower in front than behind, with slightly convex sides; posterior margin markedly medially excised, and strongly concave. With head in full-face view, eyes large but less than maximum width of antennal scape, located more to the front and not splitting the sidelines of head. Scape subcylindrical, with rounded leading edge, not reaching the posterior margin of head. Median lobe of clypeus not projecting anteriorly, anterior margin truncate and strongly notched medially. Mandible triangular, the masticatory margin bearing eight short, robust, and distinct teeth. With mesosoma in profile, the outline of dorsum a continuous convexity, with rounded junction of propodeal dorsum and declivitous surface. Mesopleural sulcus not clearly visible or absent. Hind legs with rounded basitarsus, the inside surface of which is not concave. In dorsal view, petiole node anteriorly rounded and posteriorly truncate. Sculpture of head dorsum finely ruguloreticulate and usually superimposed with punctures which become larger on the front toward the posterior margin; the lateral surface reticulate-punctulate. Mandibles faintly rugulose basally, with scattered piligerous punctures, and increasingly smooth and shining approaching the apical margin; in some specimens mandibles striate or smooth and shiny apart from the piligerous pits. Dorsum of mesosoma and petiole, and first two gastral tergites, with sparse and coarse shallow punctures, the spaces between which are smooth or with closely spaced, small punctures. Lateral portion of mesosoma and petiole node densely and finely reticulate to reticulate rugulose, superimposed with small punctures; occasionally the petiole is coarsely punctate or with effaced large punctures. Brown-yellowish, slender and erect hairs covering the entire body except the lateral portion of mesosoma; pubescence quite abundant. Integument matte or shiny, coloration dark red to black, with lighter tip of gaster and appendages. Queen. Measurments (n=7): HL: 2.52-2.81, HW: 2.44-2.81, CI: 96-101, SL: 1.86-2.07, SI: 69-76, EL: 0.48-0.51, OI: 17-20, PW: 2.00-2.31, WL: 3.82-4.16, NL: 1-1.11, NW: 1.34-1.50, NH: 1.41-1.51, DNI: 131-142, LNI: 136-143. The queen of P. perroti is relatively similar to workers, but with the typical differences of the queen caste: Head relatively broader, body size much larger, and mesopleural sulcus distinct. Discussion: Pachycondyla perroti is recognized by the following combination of characters: continuous convexity of the dorsal outline of its mesosoma, nearly rounded junction of the propodeal dorsum and declivity, strong median excision of the posterior cephalic margin, and strongly concave anterior margin of clypeus. Along its geographical range, P. perroti shows a large range of phenotypic variation. Based on these morphological variations, and the scarcity of specimens collected earlier in Madagascar, this species likely misled taxonomists, who erected a separate subspecies, P. perroti admista Forel, for a different populations. Forel (1892) had to decide on the species-level status of admista without being able to compare it with the type of P. perroti or with additional samples of P. perroti throughout its range. However, the large number of samples of P. perroti collected during recent ant inventories in Madagascar, covering most of this ant���s distributional range, present evidence to synonymize the subspecies perroti admista under P. perroti. Distribution and biology: Pachycondyla perroti is endemic to Madagascar, generally occurs in the humid habitats in the east of the island, ranging from the littoral region to the mountaintops, as well as the transitional forests of the Ampasindava peninsula (Ambilanivy Forest) and Daraina in the western slope of the northern part of the island (Fig. 60). This species is also known to occupy secondary and disturbed habitats. Although the large size of the species offers an opportunity for behavioral studies, very little is known about its biology. Field work over the past 15 years has found this species foraging most frequently on the ground and in leaf litter, and very rarely on lower vegetation. It usually nests in rotten logs, soil layers, or rarely tree stumps. Other material examined: MADAGASCAR: Antsiranana: Ambohitsara, 10 km SW Antalaha (MCZC); Chutes de la Mort (E.S. Ross) (MCZC); 10 km Cap Est, 5 km W, -15.36667, 50.43333, 20 m, lowland secondary forest (B.L. Fisher) (MCZC); 14 km W Cap Est, Ambato, -15.29128, 50.33803, 100 m, secondary rainforest (G.D. Alpert) (MCZC); 2.0 km S Andrakata, -14.65, 49.71667, 520 m, disturbed rainforest (B.L. Fisher) (CASC); 30 km N Antalaha, 3 km W of a hill (G.D. Alpert) (MCZC); 5 km SW Antalaha, -14.93806, 50.26167, 50 m, secondary forest (G.D.Alpert) (MCZC); 5 km SW Antalaha, secondary forest (G.D. Alpert) (MCZC); RS Anjanaharibe-Sud, 6.5 km SSW Befingotra, -14.75, 49.5, 875 m, rainforest (B.L. Fisher) (CASC); Ampasindava, F d'Ambilanivy, 3.9 km 181�� S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (CASC); Betaolana Forest, Ambodihazovolabe village along Ambolokopatrika River, -14.54484, 49.45163, 740 m, disturbed forest patch next to tavy (B.L. Fisher et al.) (CASC); F Ambanitaza, 26.1 km 347�� Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (CASC); F d' Antsahabe, 11.4 km 275�� W Daraina, -13.21167, 49.55667, 550 m, tropical dry forest (B.L. Fisher et al.) (CASC); Fotodriana, Cap Masoala, -15.69694, 50.27028, 25 m, rainforest (G.D. Alpert) (MCZC); Marojejy RNI. #12, -14.44533, 49.78564, 375 m, rainforest (G.D. Alpert) (MCZC); PN Marojejy, Manantenina River, 27.6 km 35�� NE Andapa, 9.6 km 327�� NNW Manantenina, -14.435, 49.76, 775 m, rainforest (B.L. Fisher) (CASC); PN Marojejy, Manantenina River, 28.0 km 38�� NE Andapa, 8.2 km 333�� NNW Manantenina, -14.43667, 49.775, 450 m, rainforest (B.L. Fisher) (CASC); PN Marojejy, -14.43817, 49.774, 488 m, rainforest (Rin'Ha, Irwin) (CASC); PN Montagne d'Ambre, 3.6 km 235�� SW Joffreville, -12.53444, 49.1795, 925 m, montane rainforest (Fisher, Griswold et al.) (CASC), PN Montagne d'Ambre, Antomboka, -12.51269, 9.17807, 970 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Manongarivo, 10.8 km 229�� SW Antanambao, - 13.96167, 48.43333, 400 m, rainforest (B.L. Fisher) (CASC); RS Manongarivo, 12.8 km 228�� SW Antanambao, - 13.97667, 48.42333, 780 m, rainforest (B.L. Fisher) (CASC); Fianarantsoa: F d'Ambalagoavy Nord, Ikongo, Ambatombe, -21.8275, 47.33889, 625 m (R. Harin'Hala & M.E. Irwin) (CASC); PN Ranomafana, Miaranony Forest, 700 m, montane forest (E. Rajeriarison) (MCZC); Mahajanga: RS Marotandrano, Marotandrano 48.3 km S Mandritsara, -16.28322, 48.81443, 865 m, transition humid forest (B.L. Fisher et al.) (CASC); Toamasina: 19 km ESE Maroantsetra, -15.48333, 49.9, 350 m, rainforest (P.S.Ward) (MCZC); 5.3 km SSE Ambanizana, Andranobe, - 15.66667, 49.96667, 600 m, rainforest (B.L. Fisher) (CASC); 6.3 km S Ambanizana, Andranobe, -15.6813, 49.958, 100 m, rainforest (B.L. Fisher) (CASC); Ile Sainte Marie, F Kalalao, 9.9 km 34�� Ambodifotatra, -16.9225, 49.88733, 100 m, rainforest (B.L. Fisher et al.) (CASC); Mahavelona (Foulpointe); -17.66667, 49.5, sandy forest (A. Pauly) (CASC); Montagne d'Akirindro 7.6 km 341�� NNW Ambinanitelo, -15.28833, 49.54833, 600 m, rainforest (Fisher, Griswold et al.) (CASC); Montagne d'Anjanaharibe, 18.0 km 21�� NNE Ambinanitelo, - 15.18833, 49.615, 470 m, rainforest (Fisher, Griswold et al.) (CASC); Nosy Mangabe, 7.43 km S Maroantsetra, - 15.4973, 49.76223, 5 m, littoral rainforest edge (B.L. Fisher et al.) (CASC); PN Mananara-Nord, 7.1 km 261�� Antanambe, -16.455, 49.7875, 225 m, rainforest (B.L. Fisher et al.) (CASC); Parcelle K7 Tampolo, -17.28333, 49.41667, 10 m, littoral forest (Malagasy ant team) (CASC); R��serve. Ambodiriana, 4.8 km 306�� Manompana, along Manompana River, -16.67233, 49.70117, 125 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, Camp Rendrirendry 34.1 km 332�� Toamasina, -17.924, 49.19967, 390 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, Camp Vohitsivalana, 37.1 km 338�� Toamasina, -17.88667, 49.2025, 520 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, 35.1 km NW Toamasina, -17.91801, 49.20074, 500 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, 34.08 km 332�� Toamasina, -17.91977, 49.20039, 525 m, rainforest (B.L. Fisher) (CASC); RNI Betampona, 34.1 km 332�� Toamasina, -17.916135, 49.20185, 550 m, rainforest (B.L. Fisher) (CASC); RS Ambatovaky, Sandrangato River, -16.81753, 49.29498, 360 m, rainforest (B.L. Fisher et al.) (CASC); RS Ambatovaky, Sandrangato River, -16.81745, 49.2925, 400 m, rainforest (B.L. Fisher et al.) (CASC); RS Ambatovaky, Sandrangato River, -16.77274, 49.26551, 450 m, rainforest (B.L. Fisher et al.) (CASC); RS Ambatovaky, Sandrangato River, -16.76912, 49.26704, 475 m, rainforest (B.L. Fisher et al.) (CASC); SF Tampolo, 10 km NNE Fenoarivo Atn, -17.2825, 49.43, 10 m, littoral rainforest (B.L. Fisher) (CASC); Sahafina Forest 11.4 km W Brickaville, -18.81445, 48.96205, 140 m, rainforest (B.L. Fisher et al.) (CASC); 1 km W Andampibe, Cap Masoala, -15.69361, 50.18139, 125 m, rainforest (G.D. Alpert) (MCZC); 4 km W Rantovato, Cap Masoala, rainforest (G.D. Alpert) (MCZC); F Ivohibe 55.6 km N Tolagnaro, -24.56167, 47.20017, 650 m, rainforest (B.L. Fisher et al.) (CASC)., Published as part of Rakotonirina, J. C. & B. L. Fisher, 2013, Revision of the Pachycondyla wasmannii - group (Hymenoptera: Formicidae) from the Malagasy region, pp. 101-141 in Zootaxa 3609 on pages 125-126, DOI: 10.11646/zootaxa.3609.2.1

Pachycondyla vohitravo Rakotonirina and Fisher, sp. nov. (Figures 3, 25, 64, 65, 66, 80) Holotype worker, Madagascar, Toamasina, Montagne d'Anjanaharibe, 18.0 km 21�� NNE Ambinanitelo, -15.1883, 49.615, 470 m, sifted litter (leaf mold, rotten wood), rainforest, 8-12 Mar 2003 (coll. Fisher, Griswold et al.), collection code BLF08002, specimen code CASENT0034499 (CASC). Paratype: one worker with same data as holotype but with specimen code CASENT0034500 (CASC). WORKER. Diagnosis: Anteromedian margin of clypeus generally convex; posterolateral margins of propodeum and petiole node with a series of sharp teeth or tubercles, dorsal face of node bluntly tuberculate; second gastral tergite reticulate punctate or densely and coarsely punctate interspersed with small punctures; first and second gastral tergites covered with abundant pubescence. Measurements (12 specimens): HW: 1.14-1.50, HL: 1.22-1.69, CI: 88-96, SL: 0.88-1.31, SI: 75-88, PW: 0.85-1.13, WL: 1.64-2.17, NH: 0.77-1.11, NL: 0.55-0.80, NW: 0.72-0.93, DNI: 115-141, LNI: 129-143. Description: Head more or less elongate to subquadrate and broadest at about midlength; the sides normally convex and gradually converging from level of eyes to base of mandibles; posterior margin slightly concave. Head capsule reticulate-rugulose, rims of rugulation high and sharp. Eyes small and protruding, maximum diameter about the same as or less than widest part of antennal scape. Antennal scape barely reaching posterior border of head. Anteromedian margin of clypeus convex, with very slight median notch. Mandibles smooth and shiny between sparse punctures; apical margin equipped with 9-12 teeth or denticles. With mesosoma in dorsal view, metanotal groove absent, pronotum generally densely punctate to reticulate-punctate interspersed with sparse blunt tubercles; propodeum reticulate-rugulose. In profile posterolateral margin of propodeum bordered with series of denticles or tubercles; mesopleural suture indistinct. With petiole in dorsal view, node roughly longer than broad; dorsum with scattered tubercles and small punctures, the spaces between which are smooth and shiny. In profile, petiolar node higher than long; lower half of posterior margin covered with a line of sharp teeth or denticles; subpetiolar process a simple anterior hooklike shape followed by small lobe posteriorly. Anterior face of first gastral segment straight, without shallow impression to lodge the petiolar node. Second gastral tergite reticulate punctate to densely punctate interspersed with small punctures. Standing hairs present and pubescence abundant on dorsum of head and body; pubescence particularly abundant on first and second gastral tergites. Head, mesosoma, and petiole node dark brown to reddish-brown in color; gaster and appendages brown or of lighter coloration. Discussion: Samples of P. vohitravo came from three isolated montane rainforest habitats, and each population shows a striking morphological divergence that might indicate several species are involved. This character variation include overall body size, the shape of the sides of the head, the presence or absence of a blunt angle on the middle of the convex anteromedial clypeal margin, and the strength of body sculpture. However, gradations in body size and sculpture also appear among populations within the distributional range of the species and we consider them to belong to a single variable species. Pachycondyla vohitravo may be confounded with P. antsiraka and P. tahary, but the presence of abundant pubescence on the first and second gastral tergites render it separable from both. Its first and second gastral segments also have dense punctures, the spaces between which are less or about the same as their maximum diameter. For P. antsiraka and P. tahary, the distance between two punctures on first and second gastral segments is twice as long as the maximum diameter of the punctures. Queen: unknown. Distribution and biology: Pachycondyla vohitravo is known to occupy the disjunct mountain tops of the RS Anjanaharibe-Sud and Makira Forest in the north east of Madagascar, Andranomay Forest in the high plateau and the complex humid forests of Ambatovy-Analamay in the central eastern of the island (Fig. 80). Workers of P. vohitravo have been collected through leaf litter sampling, but one colony was collected from a rotten log. Additional material examined: Province Antananarivo: 3 km 41�� NE Andranomay, 11.5 km 147�� SSE Anjozorobe, -18.4733, 47.96, 1300 m, montane rainforest (Fisher, Griswold et al.) (CASC); Province Antsiranana: RS Anjanaharibe-Sud, 9.2 km WSW Befingotra, -14.75, 49.4667, 1200 m, montane rainforest (B.L. Fisher) (CASC); Province Toamasina: Ambatovy, 12.4 km NE Moramanga, -18.8394, 48.3084, 1080 m, montane rainforest (B.L. Fisher et al.) (CASC); F Ambatovy, 14.3 km 57�� Moramanga, -18.85083, 48.32, 1075 m, montane rainforest, (Malagasy ant team) (CASC); Torotorofotsy,-18.8708, 48.3474, 1070 m, montane rainforest, marsh edge, (Malagasy ant team) (CASC)., Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 478-480

Pachycondyla tavaratra Rakotonirina and Fisher, sp. n. (Figures 3, 38-39, 62) Holotype worker: Antsiranana, For��t de Binara, 9.1 km 233�� SW Daraina, -13.26333, 49.60333, 650-800 m, under stone, rainforest, 4 Oct 2003 (B.L. Fisher et al.), collection code: BLF09750, specimen code: CASENT0077442 (CASC). Paratypes: 4 workers with the same data as holotype but specimen coded: CASENT0077438, CASENT0077440, CASENT0247228, CASENT0247229 (BMNH, MHNG, CASC). Worker diagnosis: Larger species (HW: 2.49-3.13 mm), antennal scape reaching the posterior margin of head; anterior margin of clypeus broadly triangular; eyes extending beyond the lateral border of head; dorsum of head and mesosoma finely striate, with sparse punctures; stout erect hairs on dorsum of mesosoma and petiole node erected relative to the body surface. Worker measurements (n=7): HL: 3.05-3.13, HW: 2.55-2.73, CI: 84-88, SL: 1.91-2.09, SI: 99-104, PW: 1.91-2.09, WL: 4.11-4.46, NL: 1.35-1.50, NW: 1.35-1.62, NH: 1.67-1.85, DNI: 100-114, LNI: 116-130. Description Worker. Head approximately longer than broad, sides convex and broadest around level of eyes, posterior cephalic margin almost straight. Eyes generally large, breaking outline of sides of the head. Scape elongate, reaching posterior margin of head. Anterior margin of clypeus projecting into triangular median lobe, which makes a blunt angle with mandibular surface when head is viewed in profile. Mandibular masticatory margins armed with eight to nine teeth and denticles. In profile, outline of dorsum of mesosoma forming an uninterrupted convexity, with rounded angle at junction of propodeal dorsum and declivitous surface; declivitous surface is very inclined anteriorly and has imperceptibly visible lateral margins. Mesopleural sulcus indistinct or absent. With petiole in lateral view, node thick, and junction of anterior face and dorsum rounded, whereas dorsum and posterior face meet at distinct angle. In dorsal view, anterior face rounded and posterior face truncate. Sculpture resembles that of P. comorensis, head covered with dense striation superimposed with scattered punctures; striation converging from posterolateral portion of head towards midline and level of eyes through frontal lobes. Mandibles coarsely costate interspersed with piligerous pits. Pronotum with dense and fine costulae, while mesonotum and propodeum are covered with transverse, dense and fine striation or microreticulation. Lateral portion of mesosoma densely, finely reticulate-rugulose or with striation in different directions, interspersed with sparse large punctures. Declivitous surface microreticulate or transversely finely striate. Lateral surface of petiole node and gastral segments microreticulate, their dorsum densely finely costulate to finely microreticulate. Dorsum of body covered with black, stout hairs which tend to be brown on the appendages; with mesosoma viewed from front, these hairs incline towards its midline and are more erect at dorsolateral margin. Slender, brownish hairs present on ventral surface of head, gaster, and coxae. Pubescence abundant on lateral portion of mesosoma, petiole node, and gastral segments, but almost absent from dorsum of propodeum, petiole node, and dorsum of first and anterior half of second gastral segments. Integument black, head and mesosoma with silky brilliance; articulations and apex of appendages reddish. As for P. comorensis, queen caste is not known for P. tavaratra, but some individual workers collected from several colonies possess single ocelli, which suggests that these workers might have assumed the reproductive role of the queen. Males are not examined in this revision. Research should be undertaken in the future to understand the colony structure and reproductive organization of the species. Discussion: Pachycondyla tavaratra is very similar to P. comorensis, but can be distinguished easily by the triangular projection of the anteromedian clypeal margin, the elongate standing erect hairs on the upper surface of the body, and the location of the eyes, which break the outline of the sides of the head. Distribution and biology: Pachycondyla tavaratra occurs in northern Madagascar, and was collected mostly from the high altitude rainforests of the PN Montagne d'Ambre and For��t de Binara (Fig. 62). Rarely has it been recorded from lower altitudes at Andavakoera and Binara forests. This species forages mostly on the ground and generally nests in rotten logs and soil layers or rarely under rocks. Its longer antennal scapes are suggestive of foraging conducted on the forest floor and through leaf litter. Other material examined: Antsiranana, PN Montagne d'Ambre, 975 m (G. D. Alpert) (MCZC); PN Montagne d'Ambre, 1000-1100 m (W.L. & D.E. Brown) (MCZC); PN Montagne d'Ambre: 3.6 km 235�� SW Joffreville, -12.53444, 49.1795, 925 m, montane rainforest (Fisher, Griswold et al.) (CASC); PN Montagne d'Ambre, Antomboka, -12.50035, 49.175, 885 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Antomboka, -12.51269, 49.17807, 970 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Cr��te, -12.58132, 49.13368, 1110 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Petit lac, -12.53664, 49.17412, 1130 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Pic Bades, -12.5186, 49.18625, 900 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Roussettes, -12.52574, 49.17238, 1025 m, montane rainforest (B.L. Fisher et al.) (CASC); F d'Andavakoera, 21.4 km 75�� ENE Ambilobe; 4.6 km 356�� N Betsiaka, -13.11833, 49.23, 425 m, rainforest (B.L. Fisher et al.) (CASC); F de Binara, 9.1 km 233�� SW Daraina, - 13.26333, 49.60333, 650-800 m, rainforest (B.L. Fisher et al.) (CASC); F de Binara, 9.4 km 235�� SW Daraina, - 13.26333, 49.6, 1100 m, montane rainforest (B.L. Fisher et al.) (CASC)., Published as part of Rakotonirina, J. C. & B. L. Fisher, 2013, Revision of the Pachycondyla wasmannii - group (Hymenoptera: Formicidae) from the Malagasy region, pp. 101-141 in Zootaxa 3609 on pages 128-129, DOI: 10.11646/zootaxa.3609.2.1

Pachycondyla comorensis (Andr��) (Figures 1, 4, 38-39, 59) Ponera comorensis Andr��, 1887: 292. Lectotype worker, present designation, Antsiranana, Nosy Be (Ernest Andr�� 1914) AntWeb specimen code: CASENT0101406 (MNHN) [examined]. Paralectotype workers with the same data but pins coded as CASENT0101407 (MNHN) and CASENT0102011 (MSNG) [examined]. [Combination in Bothroponera: Forel, 1891: 129, pl. 4, fig. 5; Dalla Torre 1893: 36; Combination in Pachycondyla (Bothroponera): Emery, 1901: 45, Forel, 1907: 14, Emery 1911: 78; Combination in Bothroponera: Wheeler, 1922: 1007; Combination in Pachycondyla by Bolton, 1995: 304]. Worker diagnosis: Larger species (HW: 2.49-3.13 mm), antennal scape extending beyond the rear cephalic border; anterior margin of clypeus truncate; eyes not breaking the outline of the sides of head; dorsum of head and mesosoma finely striate, with sparse punctures; stout erect hairs on dorsum of head and the rest of body much more inclined to the body surface. Worker measurements (n=10): HL: 2.94-3.49, HW: 2.49-3.13, CI: 85-92, SL: 2.47-2.95, SI: 94-99, PW: 1.95-2.30, WL: 4.13-4.98, NL: 1.34-1.62, NW: 1.45-1.67, NH: 1.64-1.94, DNI: 99-115, LNI: 110-126. Description: Worker. Head roughly as long as broad, broadest behind eyes on posterior third; sides very slightly convex but converging near base of mandibles; posterior margin weakly emarginate medially. Eyes moderately large, located more to the front and not breaking outline of sides of head. Antennal scape relatively long and extending beyond posterior cephalic margin. Anterior margin of clypeus truncate, not projecting into lobe, but rather straight or feebly notched medially; with head in profile, median portion perpendicular to mandibular surface. Mandibles triangular, apical margins bearing eight to nine teeth and denticles. With mesosoma in profile, outline of dorsum almost continuously convex, without distinct angle between propodeal dorsum and declivity, lateral margins of the latter also generally indiscernable. Mesopleural sulcus visible but incomplete. With petiole in dorsal view, anterior margin rounded and posterior margin straight. In profile, petiole nodiform, with rounded anterodorsal portion and distinct angle in the posterodorsal margin. Mandibles costate or sometimes smooth with effaced fine striation, and with scattered punctures from which hairs arise. Dorsum of head densely striate and superimposed with piliferous punctures from level of eyes and frontal lobes; striation radiating towards occipital corners on each side of midline of head. Dorsal sculpture of pronotum variable, either densely finely costulate or smooth and shiny with trace of striation between sparse punctures, mesonotal and propodeal dorsum with transverse, dense, and fine striation or microreticulation which becomes fairly effaced on some specimens. Lateral portion of mesosoma characterized by a mixture of fine and dense striation and reticulate���rugulation. Declivitous surface transversely finely striate or almost smooth. Lateral portion of petiole node and first two gastral segments microreticulate to densely finely striate, with scattered large punctures which turn into a smooth and shiny surface on dorsum. Upper surface of body with suberect or appressed black, stout hairs, which are shorter on the dorsum of the head and become yellowish brown to brown on appendages; with mesosoma in frontal view, these hairs inclined towards the midline of the mesosoma dorsum, suberect along the dorsolateral margin and much more appressed near the midline. Slender brownish hairs present on ventral surface of head, gaster, and coxae. Body covered with abundant pubescence except head, dorsum of propodeum, petiole node, and third and anterior half of fourth abdominal segments. Coloration is black with reddish articulations and apices of appendages. The queen caste is unknown for P. comorensis. Discussion: Pachycondyla comorensis can be easily confused with P. tavaratra at first glance. Yet the shape of the anterior clypeal margin, the location of the eyes, and the standing degrees of pilosity on the dorsum of the body allow the separation of these two species. Pachycondyla comorensis has a truncated anterior clypeal margin, eyes that do not break the outline of the sides of the head, and subdecumbent to decumbent stout hairs on the dorsum of the body. The morphological similarity of these two species could be attributed to their adaptation to different ecological habitats in northern Madagascar. Pachycondyla comorensis generally inhabits dry forest and lowland rainforest habitats, whereas P. tavaratra occupies montane rainforests. Data from different collecting events over several years suggest that these large species may not have a morphologically typical queen caste as do other species within the genus, but reproduce through one or more ergatoid or gamergates in the same colony. The absence of alate queens reduces the spatial connection between geographically distant populations because dispersal must occur by budding. Distribution and biology: Pachycondyla comorensis is known only from Madagascar, where it generally occupies dry and lowland humid forests and the coastal region in the north of the island (Fig. 59). Despite its name, P. comorensis is not known from the Comoros. This is one of two species with longer antennal scapes which are probably used to forage on the soil surface and in leaf litter. Colonies have been found frequently in the ground, in rotten logs, under stones, rarely in dead twigs and rotten sticks on the ground, and under layers of roots and litter on rock. Other material examined: MADAGASCAR: Antsiranana: [Nd. Madagascar, Amber Gebirge Nd. Madagascar H. Rolle, Berlin, S.W.11.] (MHNG); [Nossi-b��; Museum Paris Collection, Ernest Andr�� 1914] (MNHN); [Nossi-b��] (MSNG); F Lokobe, Nossi-Be Island (E.S. Ross) (MCZC); Nossi-Be (G.B. King) (MCZC); Nosy Be, 4 km ESE Andoany (=Hellville), - 13.41667, 48.3, 200 m, rainforest (P.S. Ward) (MCZC); Nosy Be, Lokobe Forest, -13.41639, 48.30722, 20 m, lowland forest (G.D. Alpert) (MCZC); Nosy Be: RNI Lokobe, 6.3 km 112�� ESE Hellville, -13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (CASC); Nosy Be: 4 km ESE Andoany (=Hellville), -13.41667, 48.3, 200 m, rainforest (P.S. Ward) (PSWC); NE, 80 km N Ambilobe; no. 2103 (J.M. Wilson) (BMNH); 5 km S Sambava, - 14.65, 50.16667, coastal forest and vanilla (W.L. & D.E. Brown) (MCZC); ridge behind Sambava, second growth forest (W.L. & D.E. Brown) (MCZC); Ampasindava, F d'Ambilanivy, 3.9 km 181�� S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (CASC); F Ambato, 26.6 km 33�� Ambanja, -13.4645, 48.55167, 150 m, rainforest (B.L. Fisher) (CASC); F d' Antsahabe, 11.4 km 275�� W Daraina, -13.21167, 49.55667, 550 m, tropical dry forest (B.L. Fisher) (CASC); F d'Ampombofofo, -12.09949, 49.33874, 25 m, littoral forest (B.L. Fisher et al.) (CASC); F d'Ampondrabe, 26.3 km 10�� NNE Daraina, -12.97, 49.7, 175 m, tropical dry forest (B.L. Fisher et al.) (CASC); F d'Analabe, 30.0 km 72�� ENE Daraina, -13.08333, 49.90833, 30 m, littoral rainforest (B.L. Fisher et al.) (CASC); F de Bekaraoka, 6.8 km 60�� ENE Daraina, -13.16667, 49.71, 150 m, tropical dry forest (B.L. Fisher et al.) (CASC); F de Binara, 7.5 km 230�� SW Daraina, -13.255, 49.61667, 375 m, tropical dry forest (B.L. Fisher et al.) (CASC); RS Manongarivo, 10.8 km 229�� SW Antanambao, -13.96167, 48.43333, 400 m, rainforest (B.L. Fisher) (CASC); RS Manongarivo, 12.8 km 228�� SW Antanambao, -13.97667, 48.42333, 780 m, rainforest (B.L. Fisher) (CASC); RS Ankarana, 7 km SE Matsaborimanga, -12.9, 49.11667, 150 m, rainforest (P.S. Ward) (PSWC); RS Ankarana, -12.90056, 49.14722, 150 m (G.D. Alpert) (MCZC); RS Ankarana, 13.6 km 192�� SSW Anivorano Nord, -12.86361, 49.22583, 210 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Ankarana, 22.9 km 224�� SW Anivorano Nord, -12.90889, 49.10983, 80 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Ambre, 3.5 km 235�� SW Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest (Fisher, Griswold et al.) (CASC); Makirovana Forest, -14.103, 50.0198, 390 m; -14.1707, 49.9541, 415 m; -14.1604, 49.9522, 550 m, rainforest (Fisher et al.) (CASC)., Published as part of Rakotonirina, J. C. & B. L. Fisher, 2013, Revision of the Pachycondyla wasmannii - group (Hymenoptera: Formicidae) from the Malagasy region, pp. 101-141 in Zootaxa 3609 on pages 120-122, DOI: 10.11646/zootaxa.3609.2.1

Pachycondyla mialy Rakotonirina and Fisher, sp. nov. (Figures 2, 9, 14, 49, 50, 51, 76) Holotype worker: Madagascar, Fianarantsoa, PN Ranomafana, Vatoharanana River, 4.1 km 231�� SW Ranomafana, -21.29, 47.4333, 1100 m, montane rainforest, ex rotten log, 27-31 Mar 2003 (Fisher, Griswold et al.), collection code BLF08535, specimen code CASENT0497667 (CASC). Paratype: 1 worker with the same data as holotype but specimen coded as CASENT0009727 (CASC). WORKER. Diagnosis: Anterior margin of clypeus wide and straight, with median notch; eyes small, with seven to eight ommatidia; metanotal groove absent; posterior face of petiolar node with sparse pubescence and lacking numerous slender hairs; anterior face of first gastral segment not forming a shallow cavity; antennal scape and outer surface of each tibia covered with shorter, erect hairs. Measurements (2 specimens): HW: 1.15, HL: 1.38-1.41, CI: 81-83, SL: 0.94-0.95, SI: 82, PW: 0.87-0.88, WL: 1.78, NH: 0.59-0.62, NL: 0.46-0.47, NW: 0.63-0.66, DNI: 137-140, LNI: 125-134. Description: Head distinctly longer than broad, widest at mid-length; posterior margin weakly medially excised; the dorsum with dense and fine reticulate punctures; anterior third of lateral surface punctate and sparsely punctulate posteriorly. Eyes very small, with at most eight ommatidia, and situated in anterior fourth of head. Antennal scape short, not extending beyond posterior cephalic border; length of erect hairs on scape about half of widest portion of scape. Anterior clypeal margin widely transverse and straight, with feeble median excision. Mandibles smooth and shiny between sparse punctures. With mesosoma in dorsal view, metanotal groove lacking; in profile, dorsal outline of mesosoma simple, approximately a continuous line; posterolateral margin of propodeum covered with narrow lamellae, which project into a blunt angle near the mid-length at about the level of propodeal spiracle; mesopleural suture indistinct. Sculpture of promesonotum punctate; the punctures become more sparse toward the propodeum. With petiole in lateral view, node thick, posterior margin convex; in dorsal view, lateral margins rounding to a convex anterior face; posterior margin slightly excised medially; dorsum of node with scattered punctures. First gastral segment with straight anterior face; first and second gastral tergites laterally coarsely punctate or foveolate and dorsally smooth and shiny between punctures. Body covered with erect golden pilosity; pubescence abundant on head dorsum, promesonotum, tip of gaster, and appendages, but absent or sparse on the rest of body dorsum. Integument glossy, reddish orange with orange apex of gaster and appendages. Queen: unknown. Discussion: One can confound P. mialy with P. nosy but workers of the former are smaller, have smaller eyes, no metanotal groove and no mesopleural suture while those of the latter are characterized by a larger body size, larger eyes, the presence of a mesopleural suture and a metanotal groove reduced to a dotted line. Distribution and biology: This species is recorded only from an elevation of 1100 m in the PN Ranomafana. Two worker specimens were collected from rotten logs., Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 469-470

Pachycondyla cambouei (Forel) (Figures 6, 8, 10, 14-28, 44-57) Bothroponera cambouei Forel, 1891: 133, pl. 4, fig. 7 Lectotype worker, present designation, Madagascar, Imerina, Antananarivo (Cambou��) AntWeb CASENT0101770 (MHNG) [examined]. Paralectotype worker and queens with same data as lectotype but with specimen codes: CASENT0101028, CASENT0101029 (MHNG) and CASENT0101404, CASENT0101402 (MNHN) [examined]. [Combination in Pachycondyla (Bothroponera):Emery, 1901: 45, 1911: 78; Combination in Bothroponera: Wheeler, 1922: 1007; in Pachycondyla by Bolton, 1995: 303]. Worker diagnosis: With head in full-face view, antennal scape not surpassing posterior cephalic margin; dorsum of head and mesosoma not finely striate; dorsum of the body with erect slender hairs and pubescence; basal half of antennal scape rounded; dorsal outline of mesosoma not forming a continuous convexity; distinct angle discernable between propodeal dorsum and declivitous surface; petiole nodiform, not anteroposteriorly flattened; anterior half of fourth abdominal tergite (gastral tergite 2) mostly smooth and shiny between large punctures. Worker measurements (n=116): HL: 1.38-2.29, HW: 1.19-2.01, CI: 85-95, SL: 0.92-1.69, SI: 72-87, PW: 0.95-1.71, WL: 1.64-2.89, NL: 0.65-1.05, NW: 0.79-1.26, NH: 0.95-1.54, DNI: 110-137, LNI: 130-169. Description: Worker. In full-face view head widest at or slightly behind level of eyes; sides somewhat convex, but weakly converging in front of eyes; shape of posterior margin variable, ranging from straight to strongly medially excised. Compound eyes either large and moderately protruding, or small with diameter less than half of maximum diameter of scape; with head in full-face view, eyes breaking lateral margins of the head. Antennal scape subcylindrical, with rounded leading edge, and not surpassing posterior margin of head. Anterior clypeal margin truncate and either straight or slightly concave. Mandibles triangular, apical margins armed with seven to nine teeth and denticles. With mesosoma in profile, the dorsal outline not forming a continuous convexity, propodeal dorsum meeting declivity at a distinct angle; mesopleural sulcus usually indistinct and incomplete. Petiole nodiform; in lateral view, anterior face, dorsum, and posterior face meet in a rounded angle. With petiole node in dorsal view, posterior margin straight or medially excised. Hind basitarsus nearly rounded and not flattened dorsoventrally, basal half of inside surface not concave. Mandibles longitudinally striate and covered with sparse piligerous punctures. Dorsum of head finely reticulate-punctate or reticulate rugulose, superimposed by punctures or setae-bearing foveolae; sides with less defined sculpture, but finely reticulate-punctate or reticulate-rugulose, interspersed with small piligerous pits. Mesosoma and petiole node with variable sculpture, dorsally ranging from coarsely densely punctate to reticulate-rugose or irregularly punctate, interspersed with quite closely spaced smaller punctures; generally fine striations running to bottom of large punctures. For lateral portions, sculpturing varies from finely rugulose to transversely striate with scattered punctures. Propodeal declivity smooth and shiny or with granular surface. Third and fourth abdominal tergites covered with large, shallow punctures, interspaces either smooth and shiny or covered with fine, small punctures; in some specimen s the large punctures are irregularly spaced and much more accentuated. Dorsum of body covered with erect slender or suberect thick hairs, which are reduced in number or absent on the antennal scape and legs. Pubescence on the third and fourth abdominal tergites either dense, sparse, or lacking. Body colors range from ferruginous red to reddish brown through dark brown to black with lighter appendages. Queen. Measurements (n=14): HL: 1.53-2.18, HW: 1.41-2.09, CI: 89-98, EL: 0.28-0.42, OI: 17-21, SL: 1.02-1.66, SI: 72-82, PW: 1.21-1.87, WL: 2.36-3.45, NL: 0.63-1.02, NW: 0.94-1.34, NH: 0.92-1.35, DNI: 130-153, LNI: 129-146. Winged queens and ergatoids are present in cambouei and its variants. Ergatoid queens and workers look very similar, but the former have ocelli and incomplete thoracic sclerites while the body of alate queens is noticeably larger in general. Variation: Pachycondyla cambouei shows a remarkable range of phenotypic diversity. To faciliate discussion of this diversity, the morphological characters of the worker caste are divided into seven forms on the basis of the shape of the occipital corner, the posterior margin of the head, the size of the eyes, the form of the mesosoma, and the abundance of pubescence on the fourth abdominal tergites. However, there is no simple pattern to the degree of variation of these forms. Specifically, a few forms have very restricted geographic boundaries while others show a wide distribution along the humid forests of Madagascar. Each form of P. cambouei is described below: Form 1 (Figs 10, 15, 18, 22, 24-25, 53). Worker measurements (n=35 workers): HL: 1.75-2.17, HW: 1.59-1.99, CI: 88-95, SL: 1.24-1.57, SI: 75-85, PW: 1.23-1.67, WL: 2.15-2.89, NL: 0.82-1.02, NW: 0.96-1.24, NH: 1.16-1.54, DNI: 110-130, LNI: 130-157. This form corresponds to the typical P. cambouei, which is distinguished by much larger eyes, medially excised posterior cephalic margin and normal shape of head and mesosoma, medially excised posterior margin of petiole, and smooth or sparsely punctate fourth abdominal tergites. Some characters are variable, such as the smooth or shagreenate propodeal declivity, and slender erect hairs with reduced pubescence or shorter and much more robust erect hairs with more pubescence. A few larger worker specimens (HW: 1.86-1.99 mm) from the Masoala area in northeastern Madagascar are included in this form. These larger workers have less defined sculpture and are covered with elongate and slender erect hairs and less pubescence. At first glance, these differences are suggestive of a distinct species, but other specimens from various localities also show patterns of less enhanced sculpture, slender and long erect hairs, and a larger-sized body than a more typical P. cambouei specimen. Form 2 (Figs 8, 16, 26-27, 54). Worker measurements (n= 8): HL: 1.83-2.19, HW: 1.66-1.96, CI: 88-91, SL: 1.33-1.51, SI: 77-81, PW: 1.26-1.48, WL: 2.35-2.69, NL: 0.82-0.97, NW: 0.98-1.18, NH: 1.19-1.44, DNI: 114-121, LNI: 138-148. Known only from workers collected in the isolated rainforest of the RS Ambohijanahary. This variant is characterized by a backward extension into a lobe of each angle of the posterior margin of the head (Fig. 16). However, workers with more typical characters of P. cambouei are also found within a few colonies of this variant, and workers that show intermediate levels of protrusion between this form and form 3 (see below) also occur in FC Didy, FC Andriantantely and PN Mantadia. Workers with intermediate levels of protrusion are included in the latter form. Form 3 (Figs 6, 17, 28-29, 55). Worker measurements (n=25): HL: 1.92-2.22, HW: 1.72-2.03, CI: 86-93, SL: 1.40-1.58, SI: 77-82, PW: 1.29-1.55, AL: 2.37-3.00, NL: 0.91-0.99, NW: 1.04-1.17, NH: 1.3-1.43, DNI: 112-125, LNI: 140-152. This variant is distinguished by the fact that the posterolateral margin of the head at each occipital corner protrudes posteriorly into a hornlike tooth. Individual workers with intermediate degrees of expansion of the occipital lobes are known from Ivohibe and PN Mantadia, and this trait varies gradually along its geographical distribution to match the general diagnosis of typical P. cambouei (form 1). Interestingly, worker specimens in some colonies from the PN Ranomafana also present this gradual form of hornlike tooth, which for this form may be indicative of some biological meaning in terms of colony behavior. Form 4 (Figs 19, 30-31, 54). Worker measurements (n=10): HL: 1.74-1.97, HW: 1.55-1.82, CI: 89-92, SL: 1.25-1.48, SI: 79-86, PW: 1.21-1.38, WL: 2.14-2.55, NL: 0.84-0.99, NW: 0.97-1.14, NH: 1.22-1.34, DNI: 111-126, LNI: 135-151. Workers of this form are differentiated by the absence of erect hairs and the absence or great reduction in abundance of pubescence on the dorsum of the propodeum. Numerous specimens that show an intermediate abundance of erect hairs and pubescence on the propodeal dorsum also are known across the range of P. cambouei. Specimens of this form could probably be misidentified as P. masoala due to the absence of erect hairs on propodeal dorsum and less defined propodeal sculpture. However, the thickness of the petiolar node, which is about as broad as long when viewed laterally, combined with the gradient of characters, are those of a true P. cambouei. Form 5 (Figs 20, 32-33, 54). Worker measurements (n=5): HL: 2.10-2.29, HW: 1.82-2.01, CI: 87-88, SL: 1.54-1.69, SI: 83-85, PW: 1.49-1.71, WL: 2.71-2.88, NL: 0.92-1.05, NW: 1.15-1.26, NH: 1.48-1.52, DNI: 118-126, LNI: 141-163. Two different altitudes in PN Masoala in northeast Madagascar yielded a few workers (4) in which both anterodorsal angles of the pronotum are bidentate or tuberculate where they meet the cervical shield, and the ventral surface of the head near the occipital angles is distinctly compressed; the integument of the entire body is ferruginous red. At first glance this form appears to be a morphologically distinct species, but among the four workers, there is one specimen with a pronotum almost angulate in dorsal view, with less developed teeth, the head apparently not strongly flattened ventrally near the occipital corners. An additional worker collected from the same national park presents a rounded anterodorsal angle of the pronotum when viewed dorsally and rounded occipital angles with the head in profile. The presence of these intermediate characters seems to link this variant to the other forms of P. cambouei. Colonies with all castes of this form should be collected in order to investigate how this character variation relates to the biology of the population and to clarify relationships with the other forms. Form 6 (Figs 14, 21, 23, 34-35, 56). Worker measurements (n=30 workers): HL: 1.60-2.07, HW: 1.46-1.87, CI: 86-93, SL: 1.11-1.57, SI: 74-87, PW: 1.12-1.47, WL: 2.02-2.74, NL: 0.76-1.02, NW: 0.93-1.18, NH: 1.08-1.41, DNI: 112-135, LNI: 132-169. This is one of the forms most similar to typical P. cambouei. Workers can be recognized by the combination of the following characters: smaller eyes, straight posterior cephalic margin, slightly concave anterior clypeal margin, flagellar segments as long as broad, shagreened propodeal declivity, and less accentuated sculpture. Abdominal tergite 4 densely covered with appressed hairs when viewed dorsally; petiole node with lateral margins strongly converging anteriorly to form a bluntly rounded tip; posterior margin slightly concave with a weak median excision; body color reddish brown. This form is dominant in the humid, high altitude forests of southern Madagascar. Form 7 (Figs 36-37, 57). Worker measurements (n=20): HL: 1.38-1.65, HW: 1.19-1.44, CI: 85-92, SL: 0.92-1.13, SI: 72-80, PW: 0.95-1.18, WL: 1.64-2.07, NL: 0.65-0.81, NW: 0.79-1.05, NH: 0.95-1.17, DNI: 119-137, LNI: 139-152. As in form 6 but workers tend to have smaller body size, broader than long basal flagellar segments, straight anterior clypeal margin, short and robust mesotibiae, and less pubescence on abdominal tergite 4. In dorsal view, petiole node nearly subquadrate, with straight or slightly concave posterior margin and broadly rounded anterior margin; reddish brown to dark brown in color with lighter gaster and appendages. In the north of Madagascar, workers of this form are slightly sculptured and noticeably shiny, with much longer erect hairs, whereas those from the south possess finer, stronger sculpture and shorter hairs. In this variant, morphological diversity occurs and continuously intergrades into that of form 6. Form 7 dominates the rainforests in northern Madagascar but is absent from the PN Montagne d���Ambre. Discussion: A better understanding of species boundaries in Malagasy ants is made possible by recent improvements in collecting methods as well as intensive sampling across the entire region. Although morphological differences can distinguish several forms within P. cambouei, enough gradual variation in character states exists across their geographical range to prevent us from hypothesizing separate species for each form. For example, differences in the shape of the posterior margin and the posterior corner of the head, density and size of punctures, and other body sculpture characteristics are not consistent across the region. Madagascar is characterized by complex topography and habitat heterogeneity. A mountain plateau on the island creates habitats ranging from montane forest at high altitudes and dry forest on its western slopes to lowland rainforest and littoral forests on its eastern side. Continuous environmental gradients exist between each type of habitat. Across these ecological gradients and ecotones, a widespread species like P. cambouei would be expected to show some variation, especially if the suitiable habitats are isolated (Raxworthy & Nussbaum 1995; Ricklefs 2004; Wiens & Donoghue 2004; Wilm�� et al. 2006; Boumans et al. 2007; Smith et al. 2007; Vences et al. 2009). An example of isolated populations is the form 2 located in the relict montane rainforests of RS Ambohijanahary, which is nested in extensive dry habitats in the western slope of Madagascar. The area may have served as a habitat refugia during periods of climatic fluctuations (Wilm�� et al. 2006; Yoder and Heckman 2006; Wollenberg et al. 2008; Pearson and Raxworthy 2009; Vences et al., 2009). To better understand the morphological diversity of P. cambouei, especially in a historical context, we hope future studies will include molecular sequence data and other morphological data such as male characters which have proven useful in distinguishing cryptic species of ants (LaPolla et al. 2011). The use of molecular sequence data of P. cambouei will also help evaluate the biogeographical mechanisms that underlie the evolutionary diversification of taxa in Madagascar (as reviewed by Vences et al. 2009). Distribution and biology: Pachycondyla cambouei is an endemic, widespread species found from the north throughout the center and the south of Madagascar. It occupies mainly mesic forests at higher and lower altitudes, and also occurs in littoral and transitional forest habitats. This species dominates the representatives of the genus Pachycondyla across its distribution range. Two or more different forms within the species occur sympatrically in several localities; these sympatric groupings include forms 2 and 6; forms 3 and 6; forms 3 and 7; forms 6 and 7; forms 1, 4, and 7; and forms 1, 5, and 7. Although this species is generally terrestrial, a few specimens have been found foraging on low vegetation. Forms 2-7 occur generally on the mountaintops, while forms 1 and 4 also can be found in lowland forests. Most forage through leaf litter and rarely on the forest floor. Nest series were discovered for most forms, collected frequently from rotten logs, under stones, in the ground, in rotten tree stumps, and between root mat layers, but seldom in dead twigs or branches above the ground. Workers of P. cambouei often play dead when their nests are disturbed. Colonies of forms 5 and 6 have not yet been found, which suggests they may be soil dwellers. The smaller eyes, longer erect hairs, and abundant pubescence in form 6 are also suggestive of an underground life style. Other material examined: Form 1. MADAGASCAR: Antananarivo: Antananarivo (Cambou��) (MNHN); Tam Perinet; B.M. 1983-201, 27 Apr-3 May 1983 (J.S. Noyes, M.C. Day) (BMNH); Amparihibe (SB); 18; BMNH (E)2003-61, -15.0353, 49.5839 (K.A. Jackson, D. Carpenter) (BMNH); Andrangoloaka (Sikora) (MHNG); Antananarivo; Museum Paris, Grandidier 1893 (Cambou��) (MNHN); Im��rina; Museum Paris, Grandidier (Cambou��) (MNHN); Imerina (Cambou��) (MHNG); (W.M. Wheeler) (MCZC); 25 km NNE Ankazobe, -18.1, 47.18333, 1500 m, montane rainforest (P.S. Ward) (MCZC); 3 km 41�� NE Andranomay, 11.5 km 147�� SSE Anjozorobe, -18.47333, 47.96, 1300 m, montane rainforest (B.L. Fisher et al.) (CASC); Ankazobe, -18.31617, 47.11583, 1241 m, urban/garden (B.L. Fisher et al.) (CASC); Kaloy, -18.58998, 47.65102, 1423 m, disturbed montane rainforest (B.L. Fisher et al.) (CASC); RNI Sohisika, Sohisika 24.6 km NNE Ankazobe, -18.10322, 47.18692, 1464 m, gallery montane forest (B.L. Fisher et al.) (CASC); RS Ambohitantely, F d���Ambohitantely, 20.9 km 72�� NE Ankazobe, -18.22528, 47.28683, 1410 m, montane rainforest (Fisher, Griswold et al.) (CASC); RS Ambohitantely, F d���Ambohitantely, Jardin Botanique, 24.1 km 59�� NE Ankazobe, -18.17139, 47.28182, 1620 m, montane rainforest (Fisher, Griswold et al.) (CASC); Tsinjoarivo, E Ambatolampy (W.L. Brown) (MCZC); Angavokely, -18.93333, 47.75 (B. Pettersson) (PSWC); Vanjamanitra, Antananarivo, 1380 m (J.M. Betsch) (MCZC); Bongolava, Pref. Tsiroanomandidy (A. Peyrieras) (MCZC); Antsiranana: 9.2 km WSW Befingotra, RS Anjanaharibe-Sud, -14.75, 49.46667, 1200 m, montane rainforest (B.L. Fisher) (CASC); Ampasindava, F d'Ambilanivy, 3.9 km 181�� S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (CASC); Ankobahoba, 32.3 km N Ambanja, -13.39166, 48.48249, 41 m, disturbed littoral rainforest (B.L. Fisher et al.) (CASC); Betaolana Forest, along Bekona River, -14.52996, 49.44039, 880 m, rainforest (B.L. Fisher et al.) (CASC); F Ambanitaza, 26.1 km 347�� Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (CASC); F d'Anabohazo, 21.6 km 247�� WSW Maromandia, -14.30889, 47.91433, 120 m, tropical dry forest (Fisher, Griswold et al.) (CASC); F de Binara, 9.1 km 233�� SW Daraina, -13.26333, 49.60333, 650-800 m, rainforest (B.L. Fisher et al.) (CASC); F de Binara, 9.4 km 235�� SW Daraina, -13.26333, 49.6, 1100 m, montane rainforest (B.L. Fisher et al.) (CASC); F d'Orangea, 3.6 km 128�� SE Remena, -12.25889, 49.37467, 90 m, littoral rainforest (Fisher, Griswold et al.) (CASC); Marojejy R.N.I. #12, -14.44533, 49.78564, 375 m, rainforest (G.D. Alpert) (MCZC); Marojejy, tributary Manantenina R., - 14.43333, 49.75, 750 m (Quinter & Nguyen) (CASC); Nosy Be, RNI Lokobe, 6.3 km 112�� ESE Hellville, - 13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (CASC); PN Marojejy, Antranohofa, 26.6 km 31�� NNE Andapa, 10.7 km 318�� NW Manantenina, -14.44333, 49.74333, 1325 m, montane rainforest (B.L. Fisher) (CASC); PN Marojejy, Manantenina River, 27.6 km 35�� NE Andapa, 9.6 km 327�� NNW Manantenina, -14.435, 49.76, 775 m, rainforest (B.L. Fisher) (CASC); PN Marojejy, Manantenina River, 28.0 km 38�� NE Andapa, 8.2 km 333�� NNW Manantenina, -14.43667, 49.775, 450 m, rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, 3.6 km 235�� SW Joffreville, -12.53444, 49.1795, 925 m, montane rainforest (Fisher, Griswold et al.) (CASC); PN Montagne d'Ambre, Antomboka, -12.51269, 49.17807, 970 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Cr��te, -12.58132, 49.13368, 1110 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Manongarivo, 12.8 km 228�� SW Antanambao, -13.97667, 48.42333, 780 m, rainforest (B.L. Fisher) (CASC); Makirovana Forest: -14.1667, 49.95, 715 m; -14.1604, 49.9522, 550 m; -14.1707, 49.9541, 415 m, rainforest (B.L. Fisher et al.) (CASC); Fianarantsoa: 29.5 km WNW Tolanaro, Vasiha Mt., -24.93694, 46., Published as part of Rakotonirina, J. C. & B. L. Fisher, 2013, Revision of the Pachycondyla wasmannii - group (Hymenoptera: Formicidae) from the Malagasy region, pp. 101-141 in Zootaxa 3609 on pages 108-119, DOI: 10.11646/zootaxa.3609.2.1

Diagnosis of workers and queens of P. sikorae species-group In the Malagasy region, the P. sikorae-group is most similar to species in the recently revised P. wasmannii-group (by Rakotonirina & Fisher 2013). These two groups share many characters listed below but the sikorae-group can be distinguished by the presence of a lateral circular pit at the base of the mandibles (Fig. 1). In the field, verification of the circular pit in the smaller species may be difficult, but workers of the P. sikorae-group can be recognized by their slender body, shiny integument, and their immediate moving reaction when their nests are disturbed. Conversely, those of the P. wasmannii-group can be identified by their robust body, matte integument, and their ability to simulate death when disturbed. 1. Mandible subtriangular, masticatory margin with 9-12 teeth and denticles. 2. Mandible with dorsolateral pit or fovea near the base; lateral sulcus present, running from the base to the apex. 3. Palp formula 2, 2. 4. Frontal lobes with an anteriorly broadly rounded outer margin which is posteriorly and laterally compressed to form a narrow surface at about the level of the eyes. 5. Anteromedian margin of clypeus straight and medially notched in P. gorogota, P. haratsingy, P. ivolo, P. maeva, P. mialy, P. nosy, P. sikorae or triangular or broadly rounded in P. agnivo, P. antsiraka, P. daraina, P. rovana, P. tahary, P. vohitravo and P. zoro. 6. Antenna with 12 segments, the funiculus gradually increasing in width towards the apex and without a distinct club. 7. Compound eyes variable in size, rarely larger than the maximum diameter of antennal scape or greater (P. sikorae, P. agnivo) and most often smaller than approximately half of the maximum diameter of antennal scape; located roughly at the midline of the head when viewed in profile. 8. Apical portion of hind tibiae with two spurs, a large pectinate spur and an anterior spur that is much smaller and simple. 9. Pretarsal claws simple. 10. Metanotal groove strongly impressed in P. gorogota, P. maeva, and P. sikorae to feebly distinct in P. haratsingy and P. nosy, or absent in other remaining species. 11. Mesopleural sulcus usually indistinct, present as three to fourbroad, discontinuous grooves or as a single narrow sulcus. 12. Posterolateral margin of propodeum simply rounded or marginate (P. haratsingy, P. rovana, P. sikorae), bordered with lamellae (P. gorogota, P. maeva, P. daraina, P. zoro) or covered with a series of sharp teeth or denticles (P. antsiraka, P. tahary, P. vohitravo). 13. Propodeal spiracle slit-like. 14. Metapleural gland opening just above the posteroventral angle of mesosoma. 15. Petiole node usually thick, very rarely a flattened scale with slight posteromedian notch (P. gorogota, P. maeva). 16. Helcium approximately at the anteroventral angle of the first gastral segment (fourth abdominal segment). 17. First and second gastral segments with distinct constriction. 18. Stridulitrum absent. 19. Sting well developed. 20. Queen comparatively similar to worker, but with the following distinctive features: broader head, presence of ocelli, eye diameter greater than the greatest width of antennal scape; mesopleuron divided into an episternum and katepisternum by a transverse sulcus; the mesosomal flight sclerites fully developed; petiole node more or less flattened anteroposteriorly; moderately larger body size with usually more voluminous gaster and denser and more elongate pubescence. The worker caste of the P. sikorae-group in Madagascar can be identified by the combination of the following characters: presence of a pit on the lateral portion of the base of the mandibles (Fig. 1); the shape of the anterior clypeal margin either approximately straight with slight median notch (Fig. 2) or projecting into triangular lobe (Fig. 3); the slit-shaped opening of propodeal spiracles; metanotal groove either present or absent; hind legs with two tibial spurs, of which one is large and pectinate and the other smaller and simple; posterior margin of propodeum simple, lamellate or infrequently with a series of sharp teeth; thick petiolar node whose posterodorsal angle has no long spines and whose posterolateral margins occasionally have a series of sharp teeth or tubercles (Fig. 19); the frontal lobes are broadly rounded. Workers of the P. sikorae-group are superficially similar to those of the P. wasmannii-group, but the former have a lateral circular pit or fovea near the base of their mandibles, which is absent in the P. wasmannii-group. The shape of the anterior margin of the clypeus allows the division of the P. sikorae-group into two species complexes. First, the sikorae complex, which includes seven species, has a broadly transverse and roughly straight anterior clypeal margin, with slight median notch (Fig. 2); second the vohitravo complex, which also contains seven species, is characterized by a medially convex or bluntly angulate anterior margin of clypeus (Fig. 3). Synopsis of the taxonomic history of the P. sikorae species-group, Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 449-451

Pachycondyla planicornis Rakotonirina and Fisher, sp. n. (Figures 2, 5, 44-45, 61) Holotype worker: Antsiranana, Betaolana Forest, along Bekona River, -14.52996, 49.44039, 880 m, ex rotten log, rainforest, 5 March 2009 (B.L. Fisher et al.), collection code: BLF22566, specimen code: CASENT0151943 (CASC). Paratypes: 2 workers, same data as holotype but with following specimen codes: CASENT0247232 and CASENT0247233 (CASC, BMNH). Worker diagnosis: With head in full-face view, antennal scape not surpassing posterior cephalic margin; dorsum of head and mesosoma not finely striate, dorsum of body with erect slender hairs and pubescence; basal half of antennal scape dorsoventrally flattened, with thin leading edge; lateral portion of hind basitarsus flattened, the basal half of the inner surface concave; dorsal outline of mesosoma not forming a continuous convexity; junction of propodeal dorsum and declivitous surface angulate; petiolar node thick; anterior half of fourth abdominal tergite (gastral tergite 2) mostly smooth and shiny between large punctures. Worker measurements (n=7): HL: 2.09-2.25, HW: 1.93-2.07, CI: 90-92, SL: 1.42-1.49, SI: 70-75, PW: 1.38-1.50, WL: 2.86-3.02, NL: 0.84-0.98, NW: 1.05-1.23, NH: 1.10-1.22, DNI: 118-127, LNI: 118-149. Description: Worker. In frontal view, head longer than broad, broadest on posterior third, posterior margin strongly concave and medially excised, sides converging in front of eyes. Compound eyes large; with head in full-face view, located slightly to the front but still breaking outline of sides of head. Antenna with basally, dorsoventrally flattened scape, and thin leading edge; with head in full-face view, scape broad from base to apex, fairly short and not extending beyond posterior margin. Median lobe of clypeus not projecting anteriorly, anterior margin distinctly truncate and medially notched. Mandibles triangular, armed with eight distinct teeth and denticles. In lateral view, although propodeal declivity on a lower level relative to dorsum of mesosoma, dorsal outline not forming a continuous convexity, with propodeal dorsum joining declivity in distinct angle; mesopleural sulcus either visible or indistinct. Basitarsus of hind legs flattened laterally, basal half of the inner surface noticeably concave. In profile, petiole nodiform, rounded anterodorsally and at distinct angle posterodorsally. Mandibles coarsely striate with piligerous punctulae. Dorsum of head densely and finely reticulate-rugulose, interspersed with scattered punctures. Dorsal surface of mesosoma coarsely punctate or densely and finely reticulate-rugulose, superimposed with variable-sized punctures; lateral portion with compact and fine reticulaterugulation and scattered punctures; propodeal declivity generally smooth. Dorsal surface of node covered with sparse, large and shallow punctures; sides coarsely, shallowly punctate with effaced rugulation. Gastral tergite smooth and shiny between widely spaced, shallow punctures or with dense, small punctures. Brown, slender, erect hairs and pubescence present on dorsum of head and body. Integument dark brown to black, with lighter shade on appendages. Queen and male castes are unknown for this species. Discussion: Pachycondyla planicornis is very similar to P. cambouei and P. perroti, but it can be distinguished by the fact that the basal half of its antennal scape is dorsoventrally flattened and its hind tibia is laterally compressed. It is a locally rare species even though its geographical range on Madagascar is quite large. Distribution and biology: Pachycondyla planicornis is a rare species, restricted geographically to the mesic forests of eastern Madagascar (Fig. 61). It is only known from lowland forest of the PN Marojejy and Betaolana (along Bekona River) in the northeast of the island, through the sandy forest of Mahavelona (Foulpointe) in the east, to the montane rainforest of the PN Ranomafana in the southeast. Collection data indicate that foraging is carried out on the ground surface, and nests are located mainly in rotten logs. Other material examined: Antsiranana: Marojejy, dense forest (J.M. Betsch) (MCZC); PN Marojejy,-14.4382, 49.774, 487 m, rainforest (Rin'Ha, Irwin) (CASC); Makirovana Forest, -14.1707, 49.9541, 415 m, rainforest (B.L. Fisher et al.) (CASC); Fianarantsoa: Belle Vue trail, Ranomafana National Park, -21.2665, 47.42017, 1020 m, mixed tropical forest (R. Harin'Hala) (CASC); Toamasina: Mahavelona (Foulpointe); -17.66667, 49.5, in sandy forest (A. Pauly) (CASC)., Published as part of Rakotonirina, J. C. & B. L. Fisher, 2013, Revision of the Pachycondyla wasmannii - group (Hymenoptera: Formicidae) from the Malagasy region, pp. 101-141 in Zootaxa 3609 on pages 127-128, DOI: 10.11646/zootaxa.3609.2.1

Pachycondyla sikorae Forel (Figures 11, 58, 59, 60, 78) Ponera (Euponera) sikorae Forel 1891: 127. Lectotype worker, Madagascar Central (Sikora), present designation, specimen code: CASENT0101862 (MHNG) [examined]. [Accepted by Dalla Torre 1893: 42. Raised to genus and combination in Euponera (Euponera) by Emery 1901: 46, 1911: 83, Wheeler 1922: 1008. Combination in Pachycondyla by Brown in Bolton 1995: 309]. WORKER. Diagnosis: Anterior margin of clypeus broadly transverse and straight, with slight median notch; mandible striate-punctate; posterior margin of head straight or broadly rounded; antennal scape surpassing posterior cephalic margin; metanotal groove deeply impressed; promesonotum clearly convex; posterior margin of propodeum and petiole node rounded in profile, but truncate when viewed dorsally; anterior surface of third abdominal segment straight, not forming shallow cavity; integument covered with brownish slender hairs; larger species (HW: 1.63-1.8 mm); overall a smooth and glossy black ant. Measurements (19 specimens): HW: 1.63-1.98, HL: 1.98-2.29, CI: 80-86, SL: 1.56-1.78, SI: 90-101, PW: 1.33-1.55, WL: 2.85-3.29, NL: 0.84-1.03, NW: 0.92-1.17, NH: 0.98-1.15, DNI: 105-118, LNI: 104-118. Description: Head longer than broad, lateral borders convex from posterior margin to the level of eyes, then straight towards the base of mandibles; posterior margin broadly convex; head dorsum smooth and shiny between dense and very small punctures. Eyes large and strongly protruding from head capsule; with head in full-face view, eyes situated in anterior third of lateral cephalic margins. Antennal scape long, extending over posterior border of head; three most apical antennal segments almost the same width. Anterior margin of clypeus roughly straight and weakly medially notched. Mandibles striate, interspersed with punctures; outer margin gradually converging toward midline; masticatory margins with eight teeth or denticles. With mesosoma in dorsal view, metanotal groove deeply impressed; in lateral view, dorsal outline of mesosoma not a continuous line, promesonotum and propodeum clearly convex and separated by a groove; declivitous surface convex, meeting the sides of propodeum at a blunt angle; lamellate membrane lacking on posterior margin of propodeum. Mesosoma dorsum smooth and shiny; mesopleuron, metapleuron, and anterior portion of the sides of propodeum with dense and small punctures. Petiole nodiform; about as long as broad in profile; subpetiolar process at a posteriorly acute angle; in dorsal view anterior and posterior margins convex. Anterior face of first gastral segment straight, without shallow impression to lodge the petiole node. Gastral tergites generally smooth and shiny dorsally and covered with dense and fine punctures laterally. Standing hairs slender and reduced in number, but frost-like or golden pubescence is abundant especially on appendages and lower portion of the sides of body. Body color black, with bluish reflection; appendages reddishblack and tip of gaster light brown. Discussion: Of the several species with a deeply impressed metanotal groove and straight anterior clypeal margin, P. sikorae is the largest in size (HW 1.63-1.98, WL 2.85-3.29) in the group and most easily recognized by its smooth and shiny black integument, with bluish reflection, which is covered with golden pubescence. Several winged queens of this species have been recorded across its distributional range. These queens are characterized by the presence of three ocelli, flight thoracic sclerites with shallowly impressed metanotal groove, and the presence of much denser erect hairs and pubescence on the body dorsum and appendages. Apart from these typical forms, their size looks apparently similar to that of the workers. Two colonies, collected from the PN Zahamena contained gamergates, which may function as a secondary reproductive (Peeters, pers. comm.). Distribution and biology: Pachycondyla sikorae is an endemic, widespread species that is distributed across the humid forest habitats in eastern Madagascar and occurs at a wide range of altitudes, from lowlands to high mountains. In the north of the island, its distribution ranges from the RS Manongarivo in the west to the PN Marojejy in the east, down to the PN Zahamena in the central east, through the PN Ranomafana and down to the PN Andohahela in the south. It generally nests in rotten logs and dead branches on the forest floor. Worker specimens have been found foraging mostly on the ground and in the leaf litter, and rarely on low vegetation. Additional material examined: Province Antsiranana: 9.2 km WSW Befingotra, RS Anjanaharibe-Sud, - 14.75, 49.4667, 1200 m, montane rainforest (B.L. Fisher) (CASC); Betaolana Forest, along Bekona River, -14.53, 49.4404, 880 m, rainforest (B.L. Fisher et al.) (CASC); F Ambato, 26.6 km 33�� Ambanja, -13.4645, 48.5517, 150 m, rainforest (B.L. Fisher) (CASC); Makirovana Forest, -14.1707, 49.9541, 415 m, rainforest, (B.L. Fisher et al.) (CASC); Makirovana Forest, -14.1604, 49.9522, 550 m, rainforest (B.L. Fisher et al.) (CASC); PN Marojejy, Antranohofa, 26.6 km 31�� NNE Andapa, 10.7 km 318�� NW Manantenina, -14.4433, 49.7433, 1325 m, montane rainforest (B.L. Fisher) (CASC); PN Marojejy, Manantenina River, 27.6 km 35�� NE Andapa, 9.6 km 327�� NNW Manantenina, -14.435, 49.76, 775 m, rainforest (B.L. Fisher et al.) (CASC); PN Marojejy, Manantenina River, 28.0 km 38�� NE Andapa, 8.2 km 333�� NNW Manantenina, -14.4367, 49.775, 450 m, rainforest, (B.L. Fisher et al.) (CASC); RS Manongarivo, 12.8 km 228�� SW Antanambao, -13.9767, 48.4233, 780 m, rainforest, (B.L. Fisher) (CASC); RS Analamerana, 16.7 km 123�� Anivorano-Nord, -12.8047, 49.3738, 225 m, tropical dry forest (B.L. Fisher) (CASC); SAVA Region, district of Sambava, Marojejy National Park, 5 km W of Manantenina village, 1st Camp site (Mantella), -14.4382, 49.774, 487 m, low altitude rainforest (Rin'Ha, Mike) (CASC); Province Fianarantsoa: 2 km W Andrambovato, along Tatamaly River, -21.5117, 47.41, 1075 m, montane rainforest (B.L. Fisher et al.) (CASC); 3 km W Ranomafana, near Ifandiana, -21.25, 47.4167, 950 m, rainforest (P.S. Ward) (PSWC); 43 km S Ambalavao, PN Andringitra, -22.2333, 47, 825 m, rainforest (B.L. Fisher) (CASC); 45 km S. Ambalavao, 785 m, rainforest (B.L. Fisher) (CASC); 9.0 km NE Ivohibe, -22.4267, 46.9383, 900 m, montane rainforest (B.L. Fisher) [Sylvain] (CASC); PN Ranomafana, (L. Bartolozzi, S. Tiati & C. Raharimina) (MCZC); PN Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango, -23.8352, 46.9637, 940 m, rainforest (B.L. Fisher et al.) (CASC); PN Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango, -23.8408, 46.9575, 1250 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Ranomafana, Vatoharanana River, 4.1 km 231�� SW Ranomafana, -21.29, 47.4333, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); Province Mahajanga: RS Marotandrano, Marotandrano 48.3 km S Mandritsara, -16.2832, 48.8144, 865 m, transitional humid forest (B.L. Fisher et al.) (CASC); Province Toamasina: [Madagascar central.] (MHNG); [Madagascar, Museum Paris, Grandidier 1893] (MNHN); Ambatovy, 12.4 km NE Moramanga, -18.8394, 48.3084, 1080 m, montane rainforest (B.L. Fisher et al.) (CASC); FC Didy, -18.1983, 48.5783, 960 m, rainforest (H.J. Ratsirarson) (CASC); FC Sandranantitra, -18.0483, 49.0917, 450 m, rainforest (H.J. Ratsirarson) (CASC); Manakambahiny, near Vavatenina Forest, -17.4667, 49.35, (A. Pauly) (CASC); PN Andasibe-Mantadia, F de Mantadia, 25.7 km 248�� Moramanga, -18.814, 48.4303, 1040 m, rainforest (F.N. Raharimalala, B. Blaimer) (CASC); PN Zahamena, - 17.7336, 48.7262, 950 m, rainforest (B.L. Fisher et al.) (CASC); PN Zahamena, Besaky River, -17.7524, 48.8532, 760 m, rainforest (B.L. Fisher et al.) (CASC); PN Zahamena, Onibe River, -17.7591, 48.8547, 780 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, Camp Vohitsivalana, 37.1 km 338�� Toamasina, -17.8867, 49.2025, 520 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, Betampona 35.1 km NW Toamasina, -17.918, 49.2007, 500 m, rainforest (B.L. Fisher et al.) (CASC); Province Toliara: PN Andohahela, 10 km NW Enakara, - 24.5667, 46.8167, 420 m, rainforest (B.L. Fisher) (CASC); PN Andohahela, 10 km SSW Eminiminy, -24.75, 46.7667, 830 m, rainforest (P.S. Ward) (PSWC); PN Andohahela, 13 km NW Enakara, -24.5667, 46.8167, 900 m, rainforest (B.L. Fisher) (CASC); F Ivohibe 55 km N Tolagnaro, -24.569, 47.204, 200 m, rainforest (B.L. Fisher et al.) (CASC); PN Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro, -24.7585, 46.8537, 275 m, rainforest (B.L. Fisher et al.) (CASC); PN Andohahela, Col du Sedro, 3.8 km 113�� ESE Mahamavo, 37.6 km 341�� NNW Tolagnaro, -24.7639, 46.7517, 900 m, montane rainforest (Fisher-Griswold Arthropod Team) (CASC)., Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 474-476

Pachycondyla agnivo Rakotonirina and Fisher, sp. n.(Figures 16, 28, 29, 30, 70) Holotype worker: Madagascar, Toamasina, FC Andriantantely, -18.695, 48.8133, 530 m, sifted litter, rainforest, 7-10 Dec 1998 (H.J. Ratsirarson), collection code: HJR122, specimen code: CASENT0317589 (CASC). WORKER. Diagnosis: Anteromedian margin of clypeus broadly convex; posterolateral margins of propodeum and petiolar node smooth, without sharp teeth or tubercles; anterior surface of first gastral segment concave, forming a shallow impression to lodge the posterior surface of petiole. Measurements (3 specimens): HW: 1.14-1.15, HL: 1.36-1.40, CI: 82-84, SL: 1.01-1.03, SI: 88-90, PW: 1.01-1.03, WL: 1.96-2.04, NH: 0.87-0.89, NL: 0.63-0.68, NW: 0.94-0.96, DNI: 143-150, LNI: 129-138. Description: In full face-view, head broader posteriorly; sides slightly convex; posterior margin feebly concave. Dorsum of head reticulate-punctate anteriorly and densely punctate behind from about the posterior third. Eyes large, maximum diameter nearly the same as the greatest width of antennal scape. Scape barely reaching posterior cephalic margin. Anteromedian clypeal margin convex. Mandibles sparsely punctate, the space between punctures smooth and shiny; masticatory margin armed with 12 teeth or denticles. With mesosoma in dorsal view, metanotal groove obsolete; in profile, mesopleural suture absent or indistinct; posterior magin of propodeum without a series of sharp teeth or tubercles; junction of lateral propodeal surface and declivity angulate, angles fairly emarginate. Dorsum of mesosoma with punctate sculpture. Petiole nodiform, anterodorsal angle faintly overhanging anterior margin in lateral view; subpetiolar process consisting of convex anterior face and broadly subquadrate posterior margin; in dorsal view node slightly broader than long, lateral border roughly convex and converging in a rounded line to the more or less straight anterior face; posterior margin medially broadly convex; dorsum of node sparsely punctate. Anterior face of first gastral segment with shallow cavity, which is deeper near the insertion of petiole and becomes superficial towards the anterodorsal angle. With gaster in ventral view, anterior half of first gastral sternite with impressions on both sides of helcium; in profile anteroventral portion rounded and anteriorly projecting as hooklike process (the prora). Dorsum of first two gastral tergites smooth between piligerous punctures. Dorsum of head and body covered with erect hairs; pubescence quite abundant on head dorsum and scarce or absent on the rest of body dorsum. Body color black with bluish reflection; appendages and tip of gaster brown. Queen: unknown. Discussion: Pachycondyla agnivo can be recognized easily by the medially convex anterior margin of its clypeus and the presence of a shallow impression on the anterior face of its first gastral tergite. No striking morphological variation is observed in this species despite the great distance between collection localities in the south and the center of Madagascar. The shallow impression on the anterior face of the first gastral segment matches the shape of the posterior section of the petiolar node, suggesting this hollow plays a role in lodging the posterior face of the node. Distribution and biology: Pachycondyla agnivo is an endemic species found in mid-elevation areas of eastern Madagascar. In addition to the type locality, the species occurs in the PN Mantadia in the central east and in the PN Andohahela in the southernmost portion of the island. It was collected by sampling leaf litter in the central east of Madagascar and was only recorded once from a pitfall trap in Andohahela. Additional material examined: Province Toamasina: PN Mantadia, 895 m, rainforest (H.J. Ratsirarson) (CASC); Province Toliara: PN Andohahela, Col du Sedro, 3.8 km 113�� ESE Mahamavo, 37.6 km 341�� NNW Tolagnaro, 900 m, montane rainforest (Fisher-Griswold Arthropod Team) (CASC)., Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 458-459

Pachycondyla vazimba Rakotonirina and Fisher, sp. n (Figures 12, 48-49, 63) Holotype worker: MADAGASCAR: Mahajanga, Parc National Namoroka, 16.9 km 317�� NW Vilanandro, - 16.4067, 45.31, 100 m, ex rotten log, tropical dry forest, 12-16 Nov 2002 (Fisher, Griswold et al.), collection code: BLF06686, specimen code: CASENT0486413 (CASC). Paratypes: series of 5 workers with the same data as holotype but with specimen codes CASENT0486411, CASENT0247224, CASENT0247225, CASENT0247226, CASENT0247227 (BMNH, MHNG, CASC, PBZT). Worker diagnosis: With head in full-face view, antennal scape not surpassing the posterior cephalic margin; dorsum of head and mesosoma not finely striate; basal half of antennal scape rounded; dorsum of mesosoma and petiole node with short and thin erect hairs; junction of propodeal dorsum and declivitous surface at a distinct angle; mesopleural suture absent or incomplete; antennal segments 6, 7, and 8 nearly twice as wide as long; with petiole in dorsal view, the posterior margin broadly concave; anterior half of fourth abdominal tergite (gastral tergite 2) covered with numerous and very dense small punctures between larger ones, mesosoma and petiole node with moderate such sculpture; outer surface of hind tibia usually without erect hairs; integument matte. Worker measurements (n=10): HL: 1.59-1.75, HW: 1.38-1.50, CI: 85-90, SL: 1.08-1.22, SI: 76-82, PW: 1.06-1.21, WL: 2.04-2.33, NL: 0.75-0.84, NW: 0.91-1.05, NH: 1.05-1.18, DNI: 119-132, LNI: 133-149. Description: Worker. Head longer than broad, widest immediately behind level of eyes; sides feebly convex along their length and converging in front of level of eyes; posterior margin weakly medially excised. Compound eyes small, diameter approximately less than half maximum width of scape. Antennal scape with rounded leading edge, not attaining posterior margin of head; antennal segments 6, 7, and 8 nearly twice as wide as long. Anterior margin of clypeus truncated and straight. Mandibles triangular, the apical margins armed with seven to nine teeth or denticles. In lateral view, dorsal outline of mesosoma not continuously convex, but rather roughly straight and interrupted by nearly angulate junction of propodeal dorsum to declivitous surface. Mesopleural sulcus indistinct or absent. Propodeal declivity triangularly shaped and narrower towards the dorsal surface. Basitarsus of hind legs generally rounded, without concavity on basal half of inside surface. With petiole in dorsal view, node anteriorly convex and posteriorly broadly concave. Dorsum of head densely and finely reticulate-punctate to reticulate-rugulose, and interspersed with larger punctures. Mandibles striate with scattered piliferous pits. Dorsum of mesosoma and petiole node through the fourth abdominal segment densely finely reticulate-punctate, with sparse, shallow, and larger punctures. Sides of mesosoma and node of petiole densely and finely reticulate-rugulose. Lateral portion of abdominal segments 3 and 4 finely reticulate-punctate or with dense, small punctures. Dorsum of mesosoma, petiole node, and gaster covered with slender and short hairs which are usually absent on outer surface of hind tibiae. Body color dark brown to black, with reddish orange appendages and articulations. Integument relatively matte between the large punctures. Queen. Measurements (n=5): HL: 1.63-1.91, HW: 1.52-1.79, CI: 92-94, SL: 1.13-1.48, SI: 75-86, EL: 0.30-0.35, OI: 19-22, PW: 1.32-1.51, WL: 2.45-2.90, NL: 0.70-0.89, NW: 1.05-1.18, NH: 1.02-1.23, DNI: 123-156, LNI: 131-146. Worker characters are generally duplicated in the queen caste, except the latter is only slightly larger, and the body is covered by denser and more elongate pubescence. Discussion: Worker specimens of Pachycondyla vazimba are very similar to those of P. wasmannii, but their smaller size, the indistinct mesopleural suture, the absence of erect hairs on the outer surface of the hind tibia, and the broader than long shape of antennal segments 6, 7, and 8 render P. vazimba separable from P. wasmannii. Distribution and biology: The distribution of P. vazimba is generally limited to western Madagascar (Fig. 63). This species occurs mostly in the dry forest habitats of the PN Ankarafantsika in the north through Beza-Mahafaly Forest in the south. Interestingly, along its north-south range it also can be found in the gallery forests of the PN Isalo, For��t de Mite, and Fiherenana, and in the disjunct montane rainforest of Analavelona. When nesting in rotten logs, dead branches on the ground and in the soil layer, P. vazimba forages most often in leaf litter and infrequently on the forest floor, habits related to its smaller eyes, shorter antennal scape, and shorter legs. Other material examined: Antsiranana: Nosy Be, RNI Lokobe, 6.3 km 112�� ESE Hellville, -13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (CASC); Fianarantsoa: PN Isalo, Sahanafa River, 29.2 km 351�� N Ranohira, -22.31333, 45.29167, 500 m, gallery forest (Fisher, Griswold et al.) (CASC); dry wash, 1 km E of Isalo National Park Interpretive Center, -22.62667, 45.35817, 885 m (R. Harin'Hala) (CASC); Mahajanga: Manerinerina, 76.6 km N Antsohihy, -14.10744, 48.11046, 247 m, disturbed forest (B.L. Fisher et al.) (CASC); PN Ankarafantsika, F de Tsimaloto, 18.3 km 46�� NE de Tsaramandroso, -16.22806, 47.14361, 135 m, tropical dry forest (Fisher, Griswold et al.) (CASC); R��serve d'Ankoririka, 10.6 km 13�� NE de Tsaramandroso, -16.26722, 47.04861, 210 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Namoroka, 9.8 km 300�� WNW Vilanandro, -16.46667, 45.35, 140 m (Fisher, Griswold et al.) (CASC); PN Namoroka, 17.8 km 329�� WNW Vilanandro, -16.37667, 45.32667, 100 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Namoroka, 16.9 km 317�� NW Vilanandro, -16.40667, 45.31, 100 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Bemarivo, 23.8 km 223�� SW Besalampy, -16.925, 44.36833, 30 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Baie de Baly, 12.4 km 337�� NNW Soalala, -16.01, 45.265, 10 m, tropical dry forest (Fisher, Griswold et al.) (CASC); Mahavavy River, 6.2 km 145�� SE Mitsinjo, -16.05167, 45.90833, 20 m, gallery forest (Fisher, Griswold et al.) (CASC); PN Tsingy de Bemaraha, 10.6 km ESE 123�� Antsalova, -18.70944, 44.71817, 150 m, tropical dry forest on Tsingy (Fisher-Griswold Arthropod Team) (CASC); PN Tsingy de Bemaraha, 2.5 km 62�� ENE Bekopaka, Ankidrodroa River, -19.13222, 44.81467, 100 m, tropical dry forest on Tsingy (Fisher-Griswold Arthropod Team) (CASC); PN Tsingy de Bemaraha, 3.4 km 93�� E Bekopaka, Tombeau Vazimba, -19.14194, 44.828, 50 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CASC); Toliara: Beza Mahafaly, 27 km E Betioky, -23.65, 44.63333, 135 m, tropical dry forest (B.L. Fisher) (CASC); Beza-Mahafaly, 27 km E Betioky, -23.65, 44.63333, 135 m, tropical dry forest (B.L. Fisher) (CASC); Fiherenana, -23.17694, 43.96083, 100 m, gallery forest (Frontier Project) (CASC); F de Kirindy, 15.5 km 64�� ENE Marofandilia, -20.045, 44.66222, 100 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CASC); F de Mite, 20.7 km 29�� WNW Tongobory, -23.52417, 44.12133, 75 m, gallery forest (Fisher-Griswold Arthropod Team) (CASC); Makay Mts., -21.21836, 45.3106, 510 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.22336, 45.32628, 480 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.30997, 45.12946, 590 m, dry forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.23343, 45.32913, 460 m, gallery forest with bamboo (B.L. Fisher et al.) (CASC); PN Kirindy Mite, 16.3 km 127�� SE Belo sur Mer, -20.79528, 44.147, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CASC); R��serve Beza Mahafaly, Parcel 1, -23.65833, 44.62889, 175 m, dry forest (Alpert et al.) (MCZC); R��serve Beza Mahafaly, Parcel 1, -23.65, 44.63333, 130 m, tropical dry forest (P.S. Ward) (MCZC); Sept Lacs, - 23.52833, 44.15556, 80 m, gallery forest (Frontier Project) (CASC)., Published as part of Rakotonirina, J. C. & B. L. Fisher, 2013, Revision of the Pachycondyla wasmannii - group (Hymenoptera: Formicidae) from the Malagasy region, pp. 101-141 in Zootaxa 3609 on pages 131-132, DOI: 10.11646/zootaxa.3609.2.1

Pachycondyla gorogota Rakotonirina and Fisher, sp. nov. (Figures 12, 37, 38, 39, 73) Holotype worker: Madagascar, Antsiranana, Makirovana Forest, -14.1707, 49.9541, 415 m, rainforest, sifted litter, 28-29 Apr 2011 (B.L. Fisher et al.) collection code: BLF26523, specimen code: CASENT0166009 (CASC). WORKER. Diagnosis: Anterior margin of clypeus wide and straight, slightly medially notched; mandible smooth between punctulae; eyes small and with very slightly convex surface; antennal scape not surpassing posterior cephalic margin; antennal segments wider towards its apex; metanotal groove deeply impressed; posterolateral margin of propodeum bordered with broad lamella, which project as a tubercle at about the level of the propodeal spiracle; petiolar node anteroposteriorly flattened, its posterior margin medially notched in dorsal view; anterior face of first gastral segment not forming a shallow cavity; antennal scape and outer surface of each tibia covered with erect hairs; dorsum of head near posterior margin and rest of body covered with numerous long, slender hairs. Measurements (1 specimen): HW: 1.56, HL: 1.79, CI: 87, SL: 1.34, SI: 86, PW: 1.22, WL: 2.67, NH: 1.10, NL: 0.58, NW: 0.98, DNI: 169, LNI: 189. Description: Head elongate, narrower in front than behind and widest behind level of eyes; posterior margin feebly concave; head dorsum and its lateral surfaces densely and finely reticulate-rugulose or reticulate-punctate. Eyes very slightly convex with more than 30 ommatidia and located in anterior fourth of lateral borders of head. Antennal scape short, not reaching posterior border of head. Anterior margin of clypeus widely transverse and straight, with weak median notch. Mandibles smooth and shiny apart from hair bearing punctures; masticatory margins with ten teeth or denticles. With mesosoma in dorsal view, metanotal groove deeply impressed and sinuate; in profile, dorsal outline of mesosoma complex, mesonotum distinctly convex; mesopleural suture visible; posterior margins of propodeum covered with broad lamella extending into a lobe or tooth at the level of propodeal spiracle. Mesosoma coarsely reticulate-rugose. Petiolar node anteroposteriorly flattened in profile, anterior face roughly straight and joining the dorsum at a rounded angle; posterodorsal angle feebly overhanging the slightly sinuate posterior margin of node. In dorsal view, posterior margin medially excised. Anterior face of first gastral segment straight, without shallow cavity or impression. Head and body covered with many long, thin, and erect golden hairs; pubescence abundant on head and pronotum, and quite reduced on the rest of body dorsum. Coloration dark brown, with brown apices of antenna and tip of gaster. Queen: unknown. Discussion: Pachycondyla gorogota appears to be similar to P. maeva by the presence of the impressed metanotal groove and the antero-posteriorly flattened petiole node whose posterior margin is medially notched in dorsal view. But for P. gorgota, the posterior cephalic margin is roughly straight, the posterolateral margin of the porpodeum is bordered with much broader lamella which projects into tubercle at the level of propodeal spiracle, and the antennal scape and outer surface of tibiae are covered with erect slender hairs. In contrast, P. maeva has a more concave posterior cephalic border, narrow lamellae on the posterior margin of the propodeum and its antennal scape and outer surface of tibiae lack erect slender hairs. Distribution and biology: The species is represented by only one worker specimen, which is from the lowland rainforest of Makirovana in the northeast of Madagascar (Fig. 73). The specimen was captured by leaf litter sampling., Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 463-464

Pachycondyla daraina Rakotonirina and Fisher, sp. nov. (Figures 18, 20, 34, 35, 36, 72) Holotype worker: Antsiranana, For��t de Binara, 9.4km 235�� SW Daraina, -13.2633, 49.6, 1100 m, montane rainforest, sifted litter (leaf mold, rotten wood), 5 Dec 2003 (Fisher et al.), collection code: BLF09800, specimen code: CASENT0043301 (CASC). Paratypes: 2 workers with same data as holotype but specimens coded as: CASENT0043300, CASENT0043302 (CASC). WORKER. Diagnosis: Anterior margin of clypeus broadly convex medially; anterior surface of third abdominal segment straight, not forming shallow impression to lodge posterior surface of petiole; posterolateral margins of propodeum and petiolar node without a series of sharp teeth or tubercles, lamellae may be present on posterior margin of propodeum. In dorsal view, petiole node broader, almost twice as broad as long; in profile, subpetiolar process simple with only an anterior triangular lobe. Measurements (6 specimens): HW: 1.41-1.54, HL: 1.55-1.72, CI: 88-91, SL: 1.20-1.29, SI: 84-85, PW: 1.13-1.25, WL: 2.13-2.26, NH: 0.89-0.97, NL: 0.62-0.63, NW: 0.87-0.98, DNI: 139-156, LNI: 142-155. Description: With head in full-face view, slightly longer than broad with weakly concave posterior margin and feebly convex lateral margins which converge progressively toward the base of mandibles and without obtuse angle at or strong curve from the insertion of eye. Frontal lobe anteriorly more or less broad, outer margin medially bluntly angulate. Head capsule densely and finely reticulate-punctate, superimposed with small punctures. Eye medium-sized and protruding, with its diameter approximately as large as the widest part of scape. Antennal scape weakly surpassing posterior cephalic margin. Clypeus with weak anteriorly protruding median lobe and bluntly angulate or medially convex anterior margin. Mandible smooth and shiny between sparse piliferous punctulae; masticatory margin equipped with 12 teeth or denticles. With mesosoma in dorsal view, metanotal groove indistinct; dorsum of mesosoma densely and finely reticulate-rugose, superimposed with small punctures. In profile, mesopleural suture indistinct; lower portion of mesopleuron and propodeum coarsely rugose interspersed with punctures. Posterior margin of propodeum emarginate, without series of sharp teeth or tubercles. Petiole node higher than long in profile; series of sharp teeth absent from convex posterior margin; anterodorsal angle weakly overhanging the anterior face; subpetiolar process anteriorly hook-like and posteriorly subquadrate, separated by a concavity. In dorsal view, node roughly twice as broad as long, with effaced reticulate-punctate sculptures. Anterior face of first gastral segment truncate, without cavity to lodge the posterior surface of petiolar node. Dorsum of head and mesosoma with fewer golden short erect hairs than petiolar node and gaster; pubescence profuse over body dorsum. Integument blackish dark brown with brown ferruginous appendages and tip of gaster. Queen: unknown. Distribution and biology: Pachycondyla daraina is known only from northern Madagascar, between 800 m and 1100 m elevation, in the forest habitats of Binara near Daraina (Fig. 72). Worker specimens of this species have been recorded foraging mostly under leaf litter and rarely on the forest floor. Additional material examined: Province Antsiranana: For��t de Binara, 9.1 km 233�� SW Daraina, -13.2633, 49.6033, 800 m, rainforest (B.L. Fisher) (CASC); For��t de Binara, 9.4 km 235�� SW Daraina, -13.2633, 49.6, 1100 m, montane rainforest (B.L. Fisher) (CASC). and dorsal view of body and full-face view of head, Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 461-463

Pachycondyla ivolo Rakotonirina and Fisher, sp. nov. (Figures 6, 43, 44, 45, 74) Holotype queen: Madagascar, Toamasina, Torotorofotsy, -18.8708, 48.3474, 1070 m, montane rainforest marsh edge, ground forager, 29 Mar 2004 (Malagasy ant team), collection code: BLF10720, specimen code: CASENT0050330 (CASC). QUEEN. Diagnosis: Anterior clypeal margin wide and straight, with slight median notch; posterior surface of petiolar node covered with slender hairs and abundant pubescence; anterior surface of first gastral segment forming shallow cavity; lateral section of head and petiolar node smooth and shiny, with scattered piliferous punctures. Measurements (1 specimen): HW: 1.88, HL: 1.99, CI: 94, SL: 1.50, SI: 80, PW: 1.48, WL: 2.86, NH: 0.99, NL: 0.69, NW: 1.20, DNI: 172, LNI: 143. Description: Head subrectangular, the sides broadly convex, with slightly broader posterior portion; posterior margin weakly medially excised. Head dorsum densely and finely reticulate-punctate, the sculpture becoming densely punctulate near posterior margin; lateral section generally smooth and shining between shallow, small punctures. Eyes large and protruding from the surface of head. With head in full-face view, antennal scape not surpassing posterior border of head; anterior clypeal margin widely transverse and straight, slightly medially notched. Mandibles smooth and glossy between sparse piligerous punctures; masticatory margins armed with ten teeth; outer margin of mandibles near the apical portion strongly curving toward the midline. With mesosoma in dorsal view, thoracic sclerites fully developed; in profile, mesopleural sulcus visible; posterolateral margin of propodeum without a series of sharp teeth or denticles. Lateral surface of mesosoma densely rugulose-punctate. In dorsal view, petiole node approximately twice as broad as long; lateral margins strongly converging to a straight anterior margin. Posterior face of node with a pair of glandular spots covered with dense and long pubescence; long, slender hairs are present around these glands. In lateral view, anterior margin of node inclined posteriorly; subpetiolar process subquadrate anteriorly with hook-like acute angle posteriorly; petiole mostly smooth and shiny. Anterior face of first gastral segment, from junction of tergite and sternite to anterodorsal angle, sloped strongly backwards and forming a shallow cavity which is deeper near the junction and becomes shallower at the anterodorsal angle; cavity with scattered, elongate, slender hairs. With gaster in profile, lateromedial portions apparently projecting anteriorly to form lateral shields for first gastral sternite. Petiole mostly smooth and shiny between small piligerous punctures, first two gastral tergites generally punctate. Pilosity characterized by long golden hairs which are much denser and more slender on propodeum and petiole node; pubescence abundant and particularly dense on posterior surface of node. Integument shiny with reddish brown color and lighter appendages. Discussion: Pachycondyla ivolo is only known from a single winged queen. This species is most similar to P. haratsingy based on the presence of a shallow cavity on the anterior face of the first gastral segment and the abundant pubescence on the posterior surface of the node, but can be distinguished easily by the smooth and shiny lateral surfaces of the head and petiolar node. It also can be confounded with P. agnivo, but the straight anterior clypeal margin and compressed petiole node with the posterior surface covered by abundant pubescence render P. ivolo separable from the latter species. Distribution and biology: The alate queen specimen was found foraging on the floor of a high elevation rainforest near the marsh edge of Torotorofotsy in central eastern Madagascar (Fig. 74). ivolo: lateral and dorsal view of body and full-face of head, holotype, Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 466-467

Pachycondyla sikorae was first described as a subgenus (Euponera) of Ponera by Forel 1891: 127. The new subgenus Euponera was later raised to genus (Emery, 1901:46) and by 1911, included a heterogenous group of species placed in four subgenera: Euponera, Mesoponera, Brachyponera, and Trachymesopus (Emery 1911: 79- 86). Brown in Bolton (1994) synonymized the genus as a junior synonym of Pachycondyla, which is where it remains today though molecular results suggest that Pachycondyla is paraphyletic (Ward 2011; Schmidt 2013). Detailed taxonomic history of Euponera modified from AntCat.org 2013: Euponera Forel, 1891: 126 [as subgenus of Ponera] Type-species: Ponera sikorae, by monotypy. Euponera in Ponerinae, Ponerini: Emery, 1895: 767; Emery, 1911: 79 [subtribe Pachycondylini]; Wheeler, 1915: 37; Arnold, 1915: 63; Forel, 1917: 237; Wheeler, 1922: 648; all subsequent authors. Euponera as subgenus of Ponera: Forel, 1891: 126; Emery, 1895: 767. Euponera as genus: Emery, 1901: 46; Emery, 1909: 364; Wheeler, 1910: 135; Emery, 1911: 79; Arnold, 1915: 63; Forel, 1917: 237; Gallardo, 1918: 64; Wheeler, 1922: 648; Borgmeier, 1923: 70; Clark, 1934: 30; Creighton, 1950: 44; Chapman & Capco, 1951: 63; Bernard, 1953: 189; Wheeler & Wheeler, 1985: 256; Dlussky & Fedoseeva, 1988: 78; H��lldobler & Wilson, 1990: 11. Euponera as junior synonym of Pachycondyla: Brown, in Bolton, 1994: 164; Bolton, 2003: 166. Checklist of the Malagasy P. sikorae species-group sikorae complex: gorogota Rakotonirina and Fisher, sp. n. haratsingy Rakotonirina and Fisher, sp. n. ivolo Rakotonirina and Fisher, sp. n. maeva Rakotonirina and Fisher, sp. n. mialy Rakotonirina and Fisher, sp. n. nosy Rakotonirina and Fisher, sp. n. sikorae Forel1891 vohitravo complex: agnivo Rakotonirina and Fisher, sp. n. antsiraka Rakotonirina and Fisher, sp. n. daraina Rakotonirina and Fisher, sp. n. rovana Rakotonirina and Fisher, sp. n. tahary Rakotonirina and Fisher, sp. n. vohitravo Rakotonirina and Fisher, sp. n. zoro Rakotonirina and Fisher, sp. n., Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 451-452

Pachycondyla nosy Rakotonirina and Fisher, sp. nov. (Figures 5, 15, 52, 53, 54, 76) Holotype worker: Madagascar, Antsiranana, Makirovana Forest, -14.1707, 49.9541, 415 m, 28 Apr 2011, ex rotten log (B.L. Fisher et al.) collection code: BLF26582, specimen code: CASENT0231239 (CASC). WORKER. Diagnosis: Anterior margin of clypeus wide and straight, weakly medially notched; eyes medium, with 14 ommatidia; metanotal groove an incomplete dotted line; numerous slender erect hairs lacking and pubescence reduced from posterior face of petiole node; anterior surface of third abdominal segment straight, not forming a shallow impression; antennal scape and outer surface of each tibia covered with long, erect hairs. Measurements (1 specimen): HW: 1.44, HL: 1.62, CI: 89, SL: 1.25, SI: 87, PW: 1.09, WL: 2.34, NH: 0.76, NL: 0.58, NW: 0.80, DNI: 138, LNI: 131. Description: Head rectangular, broadest at midlength; posterior cephalic margin more or less straight. Head dorsum densely and finely reticulate-punctate; lateral section generally punctate and sparsely punctulate toward lateroventral angle. Eyes medium, with 14 ommatidia, and located at anterior fourth of head when viewed from the front. Antennal scape short, not reaching posterior cephalic margin; scape with long, erect hairs roughly equal in length to its maximum diameter. Anterior clypeal margin broadly straight, with weak median notch. Mandibles sparsely punctate with a smooth and shiny surface. With mesosoma in dorsal view, metanotal groove a simple, shallowly dotted line; in lateral view, dorsal outline roughly a continuous line; mesopleural suture obsolete; posterior margin of propodeum narrowly lamellate and projecting into a blunt angle near midlength. In dorsal view, promesonotum with reticulate-punctulate sculpture close to dorsolateral angles and punctate towards the midline; propodeum punctulate. Petiole nodiform, with convex posterior margin in lateral view; in dorsal view, posterior margin slightly convex, anterior margin broadly rounded; sculpture coarsely punctate. Anterior face of first gastral segment straight, not forming a shallow cavity. First two gastral tergites rugulose punctate, with smooth and shiny spaces between coarse punctures. Erect standing hairs present; pubescence abundant on head dorsum and promesonotum and sparse on the rest of body dorsum. Body color dark red, with brown tip of gaster and appendages. QUEEN. Measurements (2 specimens): HW: 1.65-1.68, HL: 1.74-1.75, CI: 95-96, SL: 1.35-1.36, SI: 80- 83, PW: 1.35-1.38, WL: 2.56, NH: 0.80-0.88, NL: 0.55-0.60, NW: 0.86-0.89, DNI: 147-155, LNI: 144-146. Two winged queens were captured using malaise traps from the montane humid forests in the north and in the central eastern Madagascar. They have the usual characteristics of alate queens, including three ocelli, developed thoracic sclerites, more abundant standing slender hairs; much broader head, and a wider, more anteroposteriorly flattened petiolar node. Distribution and biology: Pachycondyla nosy is known from northeastern Madagascar, where a colony nest was found in a rotten log in Makirovana Forest. The species also occurs in the disjunct montane rainforests of the PN Montagne d���Ambre and the PN Andasibe (Fig. 76), where two alate queens were recorded from malaise traps. Additional material examined: Province Antsiranana: PN Montagne d'Ambre [Petit Lac road], 1125 m, montane rainforest (R. Harin'Hala) (CASC); Province Toamasina: 7 km SE PN Andasibe Headquarters, 1050 m, montane rainforest (R. Harin'Hala) (CASC)., Published as part of Rakotonirina, J. C. & Fisher, B. L., 2013, Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar, pp. 447-485 in Zootaxa 3683 (4) on pages 471-472

Pachycondyla wasmannii (Forel) (Figures 9, 11, 50-51,64) Bothroponera wasmannii Forel 1887: 383. Lectotype worker, present designation, Madagascar, Antsiranana, Nosy Be (C. Keller) AntWeb specimen code CASENT0101039 (MHNG) [examined]. Paralectotype workers, with the same data but coded as CASENT0101040, CASENT0101041 (MHNG) [examined]. [Redescription:Forel, 1891: 128, pl. 4, fig. 4; Dalla Torre, 1893: 37. Misspelled as wasmanni: Emery, 1895: 336, Forel, 1897: 188, 196 with description of queen and male castes. Combination in Ponera (Bothroponera) and additional note on queen by Emery, 1899: 267; in Pachycondyla (Bothroponera):Emery, 1901: 45, 1911: 78; Combination in Bothroponera: Wheeler, 1922: 1008; in Pachycondyla:Bolton, 1995: 311]. Worker diagnosis: With head in full-face view, antennal scape not surpassing the posterior cephalic margin; dorsum of head and mesosoma not finely striate; basal half of antennal scape rounded; dorsum of mesosoma and petiole node with short and thin erect hairs; junction of propodeal dorsum and declivitous surface at a distinct angle; mesopleural suture distinct; antennal segments 6, 7, and 8 nearly as wide as long; with petiole in dorsal view, the posterior margin straight or with weak median notch; anterior half of fourth abdominal tergite (gastral tergite 2) covered with numerous and very dense small punctures between larger ones, mesosoma and petiole node with moderate such sculpture; outer surface of hind tibia usually with erect hairs; integument matte. Worker measurements (n=12): HL: 1.96-2.37, HW: 1.72-2.21, CI: 87-94, SL: 1.32-1.78, SI: 76-83, PW: 1.28-1.79, WL: 2.50-3.36, NL: 0.94-1.20, NW: 1.07-1.48, NH: 1.27-1.64, DNI: 105-132, LNI: 124-159. Description: Worker. In full-face view, head broadest behind level of eyes, the sides slightly convex throughout their length; the posterior margin feebly concave; eyes quite large, maximum diameter nearly equal to greatest width of antennal scape. Scape subcylindrical, relatively short, not surpassing posterior cephalic margin; segments 6, 7, and 8 almost as long as broad. Anteromedian margin of clypeus truncate and either straight or very weakly emarginated medially. Mandibles triangular, masticatory margins armed with seven to eight teeth and denticles. In profile, dorsal outline of mesosoma without uninterrupted convexity; propodeal dorsum meeting declivitous surface in a distinct angle; mesopleural sulcus distinct and complete. Hind legs with rounded basitarsus. Petiole nodiform and thick, with anterior and posterior margins that in dorsal view are respectively broadly convex and straight with weak median notch. Head finely reticulate-punctate to finely ruguloreticulate, interspersed with large punctures behind level of eyes. Mandibles striate, with sparse punctures from which hairs arise. Dorsum of mesosoma, petiole node, and first gastral tergite covered with large punctures, the spaces between which are reticulate-punctate. Lateral portion of pronotum, mesopleuron, and lower half of propodeum generally finely reticulate-rugulose or with dense, fine rugulation. Second gastral tergite particularly with very dense, piligerous, small pits between large punctures. Standing erect hairs present on dorsum of the body, with very abundant pubescence; outer surface of hind tibiae covered with erect hairs. Dark brown to black in color with usually lighter appendages. Most specimens fairly matte except those collected from rainforest habitats. Queen. Measurements (n=7): HL: 2.20-2.39, HW: 2.08-2.24, CI: 92-95, SL: 1.62-1.82, SI: 77-84, EL: 0.39-0.41, OI: 17-19, PW: 1.63-1.87, WL: 3.31-3.71, NL: 0.99-1.16, NW: 1.34-1.48, NH: 1.31-1.51, DNI: 127-148, LNI: 125-136. Winged queens are very similar to workers, but have a body modified with the general characteristics of the queen caste. Winged queen has a much broader head. Ergatoid queens also present in P. wasmannii, which look comparable to worker caste but have one ocellus and reduced thoracic sclerites. Discussion: Although P. wasmannii is very similar to P. vazimba, it can be distinguished readily by its larger size, with a distinct and complete mesopleural suture, antennal segments 6, 7, and 8 nearly as wide as long, and more erect hairs on the dorsum of mesosoma and petiole node. Apparently there are two forms found within this species. These are geographically isolated due to the presence of high mountain chains in northwestern Madagascar. The first form, in which gastral tergite 2 is densely, finely punctate between larger punctures and has a matte integument, occupies the west and southwest regions of Madagascar. In contrast, the second form is characterized by worker specimens that have a shining integument, and gastral tergite 2 is covered with nearly effaced, closely spaced, small punctures between larger, shallow punctures. This form has been collected in the northwest and northeast of the island. However, workers with intermediate degrees of these phenotypic variations also occur. Distribution and biology: Pachycondyla wasmannii is an endemic, widespread species of the Malagasy region. It occurs generally in western Madagascar and in Anjouan of the Comoros Islands (Fig. 64). Most frequently this species has been collected from the dry forests and woodland habitats in the west of Madagascar. However, it is known also from subhumid gallery forests in the western slopes, littoral, transitional, and humid forests in the north and southeast of the island, and from disjunct montane rainforests on the high plateau. This species is also capable of colonizing human-modified habitats. There are two morphological forms; the first occurs in the dry and gallery forest habitats in the west of Madagascar, whereas the second is generally known from the transitional mesic forests and some of the rainforest sites in the north and northeast of the island. Across its distribution range, P. wasmannii dominates the Pachycondyla in dry forest habitats where it is sympatric only with P. vazimba. In its remaining distribution range, other species within the genus co-occur with P. wasmannii, and P. cambouei, which is widely distributed throughout the humid forests, becomes the dominant species. Pachycondyla wasmannii nests mainly in rotten logs; however, it can also be found on rotten sticks and branches on the ground, in soil layers, under rocks, beneath litter moss on rocks, and under rotten logs. It typically forages on the forest floor and through leaf litter, and very rarely on low-growing vegetation. Workers of this species usually fake death (thanatosis) after a nest disturbance. One interesting geographic point about P. wasmannii is its presence on Anjouan Island in the Comoros but not Mayotte Island, although the latter is geographically closer to Madagascar. How it arrived on Comoros could shed light on our understanding of the historical biogeography and the origins of the Malagasy species of Pachycondyla. Other material examined: THE COMOROS: Anjouan, Mount Ntringui, -12.19865, 44.41866, 740 m, montane forest (B.L. Fisher et al.) (CASC); Mount Ntringui, -12.19641, 44.41791, 550 m, rainforest (B.L. Fisher et al.) (CASC). MADAGASCAR: Antananarivo: RS Ambohitantely, F d Ambohitantely, 20.9 km 72�� NE d Ankazobe, -18.22528, 47.28683, 1410 m, montane rainforest (Fisher, Griswold et al.) (CASC); Tsinjoarivo, E. Ambatolampy, dry forest (W.L. Brown) (MCZC), Antsiranana: [Nosib��] (V��ltzkow) (MHNG); [Nossi b�� p. Madagascar] (C. Keller) (MHNG); 14 km W Cap Est, Ambato, -15.29139, 50.33806, 200 m, secondary raiforest (Alpert et al.) (MCZC); 15 km S Sambava, 10 m, coastal forest (J.M. Betsch) (MCZC); 48 km ENE Morondava, -20.06667, 44.65, tropical dry forest (D.M. Olson) (MCZC); 5 km S Antalaha, Ambodikofo Hill (G.D. Alpert) (MCZC); 5 km SW Antalaha, -14.93806, 50.26167, 50 m, secondary forest (G.D. Alpert) (MCZC); 84 km SW Sambava to Andapa, 70-160 m, degraded forest (W.L. and D.E. Brown) (MCZC); Ambanja, -13.68268, 48.45245, 30 m, urban/garden (B.L. Fisher et al.) (CASC); Ambondrobe, 41.1 km 175�� Vohemar, -13.71533, 50.10167, 10 m, littoral rainforest (B.L. Fisher) (CASC); Ampasindava, F d'Ambilanivy, 3.9 km 181�� S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (CASC); Ankarana, -12.9, 49.1, 100 m (G.D. Alpert) (MCZC); F Ambanitaza, 26.1 km 347�� Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (CASC); F Ambato, 26.6 km 33�� Ambanja, - 13.4645, 48.55167, 150 m, rainforest (B.L. Fisher) (CASC); F d' Andavakoera, 21.4 km 75�� ENE Ambilobe; 4.6 km 356�� N Betsiaka, -13.11833, 49.23, 425 m, rainforest (B.L. Fisher) (CASC); F d' Antsahabe, 11.4 km 275�� W Daraina, -13.21167, 49.55667, 550 m, tropical dry forest (B.L. Fisher et al.) (CASC); F d'Ampombofofo, - 12.09949, 49.33874, 25 m, littoral forest (B.L. Fisher et al.) (CASC); F d'Ampondrabe, 26.3 km 10�� NNE Daraina, -12.97, 49.7, 175 m, tropical dry forest (B.L. Fisher) (CASC); PN Sahamalaza, F d'Anabohazo, 21.6 km 247�� WSW Maromandia, -14.30889, 47.91433, 120 m, tropical dry forest (Fisher, Griswold et al.) (CASC); F d'Analabe, 30.0 km 72�� ENE Daraina, -13.08333, 49.90833, 30 m, littoral rainforest (B.L. Fisher) (CASC); F de Bekaraoka, 6.8 km 60�� ENE Daraina, -13.16667, 49.71, 150 m, tropical dry forest (B.L. Fisher et al.) (CASC); F de Binara, 7.5 km 230�� SW Daraina, -13.255, 49.61667, 375 m, tropical dry forest (B.L. Fisher) (CASC); F de Binara, 9.1 km 233�� SW Daraina, -13.26333, 49.60333, 650-800 m, rainforest (B.L. Fisher et al.) (CASC); F d'Orangea, 3.6 km 128�� SE Remena, -12.25889, 49.37467, 90 m, littoral rainforest (Fisher, Griswold et al.) CASC; Marojejy RNI #12, -14.44533, 49.78564, 375 m, rainforest (G.D. Alpert) (MCZC); Montagne des Fran��ais, 7.2 km 142�� SE Antsiranana (=Diego Suarez), -12.32278, 49.33817, 180 m, tropical dry forest (Alpert et al.) (CASC); Nosy Be, 4 km ESE Andoany (=Hellville), -13.41667, 48.3, 200 m, rainforest (P.S. Ward) (PSWC); Nosy Be, Lokobe Forest, - 13.41639, 48.30722, 20 m (G.D. Alpert) (MCZC); Nosy Be, RNI Lokobe, 6.3 km 112�� ESE Hellville, -13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (CASC); PN Marojejy, Manantenina River, 28.0 km 38�� NE Andapa, 8.2 km 333�� NNW Manantenina, -14.43667, 49.775, 450 m, rainforest (B.L. Fisher) (CASC); PN Montagne d'Ambre, 3.6 km 235�� SW Joffreville, -12.53444, 49.1795, 925 m, montane rainforest (Fisher, Griswold et al.) (CASC); PN Montagne d'Ambre, Antomboka, -12.50035, 49.175, 885 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Antomboka, -12.51269, 49.17807, 970 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Pic Bades, -12.5186, 49.18625, 900 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Manongarivo, 10.8 km 229�� SW Antanambao, -13.96167, 48.43333, 400 m, rainforest (B.L. Fisher) (CASC); RS Manongarivo, 12.8 km 228�� SW Antanambao, -13.97667, 48.42333, 780 m, rainforest (B.L. Fisher) (CASC); RS Manongarivo, 14.5 km 220�� SW Antanambao, -13.99833, 48.42833, 1175 m, montane rainforest (B.L. Fisher) (CASC); RS Ankarana, 7 km SE Matsaborimanga, -12.9, 49.11667, 150 m, rainforest (P.S. Ward) (MCZC); RS Analamerana, 16.7 km 123�� Anivorano-Nord, -12.80467, 49.37383, 225 m, tropical dry forest (B.L. Fisher) (CASC); RS Analamerana, 28.4 km 99�� Anivorano-Nord, -12.74667, 49.49483, 60 m, tropical dry forest (B.L. Fisher) (CASC); RS Ankarana, -12.90056, 49.14722, 150 m, dry forest (Alpert et al.) (MCZC); RS d'Ambre, 3.5 km 235�� SW Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Ankarana, 13.6 km 192�� SSW Anivorano Nord, -12.86361, 49.22583, 210 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Ankarana, 22.9 km 224�� SW Anivorano Nord, -12.90889,49.10983, 80 m, tropical dry forest (Alpert et al.) (CASC); PN Marojejy, 8 km NW Manantenina, -14.43333, 49.78333, 450 m, montane rainforest (E.L. Quinter) (CASC); Vohemar, -13.37723, 50.0205, 25 m, cultivated land (B.L. Fisher et al.) (CASC); Makirovana Forest: -14.1604, 49.9522, 550 m, rainforest (B.L. Fisher et al.) (CASC); Makirovana Forest, - 14.17066, 49.95409, 415 m and 225 m, rainforest (B.L. Fisher et al.) (CASC); Makirovana Forest, -14.10295, 50.01984, 390 m, rainforest (B.L. Fisher et al.) (CASC); Fianarantsoa: 15 km E Sakaraha, -22.9, 44.68333, 760 m, tropical dry forest (P.S. Ward) (MCZC); 30 km NNW Ranohira, Isalo Nat. Park, -22.31722, 45.29333, 455 m, canopy forest (G.D. Alpert) (MCZC); F d'Analalava, 29.6 km 280�� W Ranohira, -22.59167, 45.12833, 700 m, tropical dry forest (Fisher, Griswold et al.) (CASC); F d'Atsirakambiaty, 7.6 km 285�� WNW Itremo, -20.59333, 46.56333, 1550 m, montane rainforest (Fisher, Griswold et al.) (CASC); PN Isalo, Ranohira, Canyon de Sinze, - 22.48333, 45.55, 800 m, rainforest (E. Rajeriarison) (MCZC); PN Ranomafana, -21.01667, 47.01667, 700 m, montane forest (E. Rajeriarison) (MCZC); PN Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango, -23.83517, 46.96367, 940 m, rainforest (B.L. Fisher et al.) (CASC); PN Isalo, 9.1 km 354�� N Ranohira, - 22.48167, 45.46167, 725 m, gallery forest (Fisher, Griswold et al.) CASC; PN Isalo, Ambovo Springs, 29.3 km 4�� N Ranohira, -22.29833, 45.35167, 990 m, Uapaca woodland (Fisher, Griswold et al.) (CASC); PN Isalo, Sahanafa River, 29.2 km 351�� N Ranohira, -22.31333, 45.29167, 500 m, gallery forest (Fisher, Griswold et al.) (CASC); 9 km NNW Ranohira, PN Isalo, -22.48333, 45.38333, 800 m, rainforest (P.S. Ward) (MCZC); Tsaranoro, 32.8 km 229�� Ambalavao, -22.08483, 46.77633, 950 m, rainforest (B.L. Fisher et al.) (CASC); Mahajanga: 124 km SE Mahajanga (G.D. Alpert) (MCZC); Ampamakiambato, 45 km SW Ambanja, -13.97545, 48.15929, 145 m, disturbed forest in tsingy (B.L. Fisher et al.) (CASC); F Ambohimanga, 26.1 km 314�� Mampikony, -15.96267, 47.43817, 250 m, tropical dry forest (B.L. Fisher) (CASC); F de Tsimembo, 11.0 km 346�� NNW Soatana, - 18.99528, 44.4435, 50 m, tropical dry forest (Fisher, Griswold Arthropod Team) (CASC); F de Tsimembo, 8.7 km 336�� NNW Soatana, -19.02139, 44.44067, 20 m, tropical dry forest (Fisher, Griswold Arthropod Team) (CASC); Mahavavy River, 6.2 km 145�� SE Mitsinjo, -16.05167, 45.90833, 20 m, gallery forest (Fisher, Griswold et al.) (CASC); Manerinerina, 76.6 km N Antsohihy, -14.10744, 48.11046, 247 m, disturbed forest (B.L. Fisher et al.) (CASC); PN Ankarafantsika, SF Ampijoroa, 5.4 km 331�� NW Andranofasika, -16.29889, 46.813, 70 m, tropical dry forest (Fisher, Griswold et al.) CASC; PN Ankarafantsika, Ampijoroa SF, 40 km 306�� NW Andranofasika, - 16.32083, 46.81067, 130 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Ankarafantsika, F de Tsimaloto, 18.3 km 46�� NE de Tsaramandroso, -16.22806, 47.14361, 135 m, tropical dry forest (Fisher, Griswold et al.) CASC; PN Baie de Baly, 12.4 km 337�� NNW Soalala, -16.01, 45.265, 10 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Namoroka, 16.9 km 317�� NW Vilanandro, -16.40667, 45.31, 100 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Namoroka, 17.8 km 329�� WNW Vilanandro, -16.37667, 45.32667, 100 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Namoroka, 9.8 km 300�� WNW Vilanandro, -16.46667, 45.35, 140 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Tsingy de Bemaraha, 10.6 km ESE 123�� Antsalova, -18.70944, 44.71817, 150 m, tropical dry forest on tsingy (Fisher-Griswold Arthropod Team) (CASC); PN Tsingy de Bemaraha, 2.5 km 62�� ENE Bekopaka, Ankidrodroa River,-19.13222, 44.81467, 100 m, tropical dry forest on tsingy (Fisher-Griswold Arthropod Team) (CASC); PN Tsingy de Bemaraha, 3.4 km 93�� E Bekopaka, Tombeau Vazimba, -19.14194, 44.828, 50 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CASC); R��serve d'Ankoririka, 10.6 km 13�� NE de Tsaramandroso, -16.26722, 47.04861, 210 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RF Beanka, 50.2 km E Maintirano, -18.02649, 44.05051, 250 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CASC); RF Beanka, 52.7 km E Maintirano, -18.0622, 44.52587, 300 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CASC); RF Beanka, 53.6 km E Maintirano, -18.04014, 44.53394, 272 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CASC); RF Beanka, 54.3 km E Maintirano, -18.06009, 44.54086, 262 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CASC); RS Bemarivo, 23.8 km 223�� SW Besalampy, - 16.925, 44.36833, 30 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Marotandrano, Marotandrano 48.3 km S Mandritsara, -16.28322, 48.81443, 865 m, transitional humid forest (B.L. Fisher et al.) (CASC); Toamasina: 1 km W Andampibe, Cap Masoala, -15.69361, 50.18139, 125 m, rainforest (G.D. Alpert) (MCZC); 8 km SW Cap Est, -15.30694, 50.45417, 15 m, littoral forest (G.D. Alpert) (MCZC); Toliara: 29 km NNW Ranohira, PN Isalo, -22.31614, 45.29625, 490 m, canopy forest (G.D. Alpert) (MCZC); 48 km ENE Morondava, Kirindy, - 20.06667, 44.65, 30 m, tropical dry forest (D.M. Olson) (CASC); 48 km ENE Morondava, Kirindy Forest, - 20.07444, 44.67611, 100 m, tropical dry forest (P. Rabeson) (MCZC); 50 kms N Morondava, -20.06667, 44.58333, primary dry forest (A. Pauly) (CASC); 9 km NNW Ranohira, P.N. Isalo, -22.48333, 45.38333, 800 m, rainforest (P.S. Ward) (BMNH); Andohahela, Parcel #1 versante E, 300 m, rainforest (L. Bartolozzi, S. Tiati & C. Raharimina) (MCZC); Bekonazy = 0.5 km S (n. of Morondava), forest with baobab (W.L. Brown) (MCZC); Beza Mahafaly, 27 km E Betioky, -23.65, 44.63333, 135 m, tropical dry forest (B.L. Fisher) (CASC); Fiherenana, - 23.17694, 43.96083, 100 m, gallery forest (Frontier Project) (CASC); FC Analavelona, 29.2 km 343�� NNW Mahaboboka, -22.675, 44.19, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); FC Analavelona, 33.2 km 344�� NNW Mahaboboka, -22.64333, 44.17167, 1300 m, montane rainforest (Fisher, Griswold et al.) (CASC); F de Beroboka, 5.9 km 131�� SE Ankidranoka, -22.23306, 43.36633, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CASC); F de Kirindy, 15.5 km 64�� ENE Marofandilia, -20.06855, 44.65956667, 30 m, tropical dry forest (B.L. Fisher) (CASC); F de Mite, 20.7 km 29�� WNW Tongobory, -23.52417, 44.12133, 75 m, gallery forest (Fisher-Griswold Arthropod Team) (CASC); F de Petriky, 12.5 km W 272�� Tolagnaro, -25.06167, 46.87, 10 m, littoral rainforest (B.L. Fisher) (CASC); F de Zombitse near Sakaraha, 650 m (E.S. Ross) (MCZC); F Ivohibe 55.0 km N Tolagnaro, -24.569, 47.204, 200 m, rainforest (B.L. Fisher et al.) (CASC); F Vohidava 88.9 km N Amboasary, -24.24067, 46.28783, 500 m, spiny forest/dry forest transition (B.L. Fisher et al.) (CASC); Makay Mts., -21.227, 45.33222, 475 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.22336, 45.32628, 480 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.22344, 45.3135, 550 m, gallery forest with bamboo (B.L. Fisher et al.) (CASC); Makay Mts., -21.31664, 45.1296, 620 m, dry forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.21836, 45.3106, 510 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.20978, 45.34184, 525 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.30997, 45.12946, 590 m, dry forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.21761, 45.33917, 500 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., - 21.23343, 45.32913, 460 m, gallery forest with bamboo (B.L. Fisher et al.) CASC); Makay Mts., -21.21985, 45.32396, 500 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Manatantely, Published as part of Rakotonirina, J. C. & B. L. Fisher, 2013, Revision of the Pachycondyla wasmannii - group (Hymenoptera: Formicidae) from the Malagasy region, pp. 101-141 in Zootaxa 3609 on pages 132-136, DOI: 10.11646/zootaxa.3609.2.1

A surprising feature of the SARS-CoV-2 pandemic to date is the low burdens reported in sub-Saharan Africa (SSA) countries relative to other global regions. Potential explanations (e.g., warmer environments 1 , younger populations 2-4 ) have yet to be framed within a comprehensive analysis accounting for factors that may offset the effects of climate and demography. Here, we synthesize factors hypothesized to shape the pace of this pandemic and its burden as it moves across SSA, encompassing demographic, comorbidity, climatic, healthcare and intervention capacity, and human mobility dimensions of risk. We find large scale diversity in probable drivers, such that outcomes are likely to be highly variable among SSA countries. While simulation shows that extensive climatic variation among SSA population centers has little effect on early outbreak trajectories, heterogeneity in connectivity is likely to play a large role in shaping the pace of viral spread. The prolonged, asynchronous outbreaks expected in weakly connected settings may result in extended stress to health systems. In addition, the observed variability in comorbidities and access to care will likely modulate the severity of infection: We show that even small shifts in the infection fatality ratio towards younger ages, which are likely in high risk settings, can eliminate the protective effect of younger populations. We highlight countries with elevated risk of 'slow pace', high burden outbreaks. Empirical data on the spatial extent of outbreaks within SSA countries, their patterns in severity over age, and the relationship between epidemic pace and health system disruptions are urgently needed to guide efforts to mitigate the high burden scenarios explored here.