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Ceropegia gilboaensis Heiduk, N.R.Crouch & D.Styles sp. nov. (Figs 3, 4 A−E, 5A) Diagnosis: —Plants of Ceropegia gilboaensis are perennial tuberous succulents with a prostrate growth habit, ovate to elliptic leaves, verrucose follicles, and a cylindrical-campanulate corolla, that can be separated from C. ngomensis (Figs 1I, 5B) in lacking the distinctly keeled or fluted corolla lobe bases that embed the gynostegium forming a characteristic 5-star shaped corolla mouth in C. ngomensis. The basally fused corolla lobes of C. gilboaensis have transverse burgundy bands but are purple-mottled in C. ngomensis. In C. gilboaensis, the inner corona lobes are adpressed to the back of the stamens along their full length and closely proximate to subconnivent over the style-head, whilst in C. ngomensis they are short and only adpressed to the base of the stamens. The outer corona lobes are deeply bifid in C. gilboaensis but undivided to form five cups with incurved margins in C. ngomensis. Type: — SOUTH AFRICA. KwaZulu-Natal, the Karkloof, Mt Gilboa, ca. 1620 m, 12 November 2020, D. G. A. Styles & A. Heiduk 5841 (holotype; NU0092539!). Description: —Perennial dwarf herb. Root a subterranean tuber, 20–40 mm in diameter, ca. 20 mm high, somewhat depressed on upper side, rounded below. Stems 1 or 2, up to 150 mm long, produced annually, branched sparingly at base, slender, creeping, prostrate, shortly pubescent. Leaves in 3–6(–8) pairs, shortly petiolate (to 1.7 mm long), lamina elliptic-ovate, 10–12 × 7–9 mm, ciliate with white hairs along the thin often purplish margins, pubescent above and below, venation prominent on lower surface and recessed on upper surface, basally rounded, apically acute. Flowers 1(–2) lateral at nodes, open in succession, with pungent fetid, slightly acidic-fecal floral scent; pedicels slender, pubescent, 6–7 mm long. Sepals narrowly triangular, 2.2–3.7 mm long, sparsely pubescent. Corolla steep cylindrical-campanulate, ca. 13–14 mm across, corolla lobes spreading, divided into fused and unfused section ± halfway; tube 4.5–6.3 mm deep, velvety, glabrous (rarely with sparse white trichomes on margins of corolla lobe tips), cream within with raised, transverse burgundy bands that proximally are more entire than distally; lobes narrowly triangular, 3.3–5.4 mm long, keeled, concave above, proximally cream with raised burgundy striations, distally velvety and brownish-red to rarely olive-green. Gynostegium sessile, glabrous, dark purple. Gynostegial corona 3.6–4.4 mm in diam., 2.0– 3.1 mm high, of staminal and interstaminal parts, interstaminal (outer) corona lobes joined to form a cup with V-shaped outer margin, deeply bifid, lobules ca. 0.3 mm long, falcate, inclined towards the inner lobes; staminal (inner) corona lobes adpressed to stamina along their entire length and incumbent on style-head, linear, ca. 1.7 × 0.3 mm, apices inclined over the staminal column, closely proximate to subconnivent. Pollinarium pollinia sagittate, ca. 450 × 400 μm, brownish, insertion crest ca. 250 μm long, stramineous; caudicles ca. 90 μm long; corpusculum sagittate, ca. 315 × 140 μm, brown. Follicle with both mericarps developing, erect, ca. 40–45 mm long and 4 mm at widest point, verrucose, green flecked with purple, maturing to purple-brown. Seeds ovate-elliptic, ca. 6 × 2 mm, brown, comose, coma ca. 7 mm, white. Additional material examined: Ceropegia gilboaensis: SOUTH AFRICA. KwaZulu-Natal, the Karkloof, Mt Gilboa, ca. 1630 m, 03 April 2022, D. G. A. Styles & A. Heiduk 6032 (NU0092540!; fruiting material). Distribution and ecology: — Ceropegia gilboaensis is to date known only from the Karkloof in the KwaZulu-Natal Midlands, South Africa. It is the only representative of the genus documented from the Karkloof, and is considered endemic there, at elevations of 1320−1670 m. This new species is found within grassland, growing exposed in shallow soil on the margins of, and in between, dolerite rock outcrops. Common grasses at the sites include Aristida junciformis Trin. et Rupr. (1842: 143) and Sporobolus natalensis (Steudel 1855: 154) Durand & Schinz (1894: 822). Associated herbaceous species include Aloe neilcrouchii Klopper & Smith (2010: 95), Crassula dependens Bolus (1881: 391) and Ledebouria sandersonii (Baker 1870: App. 5) Venter & Edwards (2003: 50). The national vegetation map shows that C. gilboaensis occurs in Midlands Mistbelt Grassland and Mooi River Highland Grassland (South African National Biodiversity Institute 2006–2018). In this area, Midlands Mistbelt Grassland appears to have been mapped as occurring below a contour approximating an elevation of 1500 m, and Mooi River Highland Grassland above this. However, this likely represents an over-simplification of the floristics in contact areas, where C. gilboaensis occurs just above 1500 m. Midlands Mistbelt Grassland is described as “forb-rich, tall, sour Themeda triandra [Forsskal (1775: 178)] grasslands transformed by the invasion of native Ngongoni grass (Aristida junciformis subsp. junciformis). Only a few patches of the original species-rich grasslands remain.” (Mucina et al. 2006: 422). This vegetation type is “Endangered (one of the most threatened vegetation types of KwaZulu-Natal)” according to Mucina et al. (2006: 423). Furthermore, the Mistbelt of KwaZulu-Natal has been identified as an important centre of endemism (Mucina et al. 2006), although not yet formally recognized as such (Mucina et al. 2006; Van Wyk & Smith 2001). More limited floristic information is provided on Mooi River Highland Grassland, which is also considered threatened, but with the status of Vulnerable rather (Mucina et al. 2006). Phenology: —Seen in flower from October to December, and in fruit from Jan uary to April. Etymology: — Ceropegia gilboaensis is named for the type locality at Mt Gilboa in the Karkloof, SouthAfrica, where it was discovered in 2008 by Isabel Johnson, a field botanist, ecological consultant and dedicated conservationist. Conservation Status: — Ceropegia gilboaensis is only known from two localities 7.5 km apart from each other. At each locality, plants were observed within an area of less than 1 km 2. However, a study of aerial imagery indicates that within the surrounding landscape there may be a further 35 km 2 of similar habitat in which C. gilboaensis could potentially occur. The total number of plants is estimated at less than 250 individuals. The grassland habitat at one of the two localities suffers from the trampling effects of cattle, but does not otherwise appear to be under anthropogenic threat. The species’ conservation status is suggested as Endangered (EN) under IUCN criteria B2(a)(b)(iii) and D (IUCN Standards and Petitions Subcommittee 2019). Discussion Ceropegia gilboaensis is a new species in section Chamaesiphon, the largest section in Ceropegia s.l. with 115 tuberous species (Bruyns et al. 2017). The overwhelming majority (ca. 90%) of these are species with open non-trapping, brood-site mimicking flowers; species with kettle-trap flowers have evolved in at least three subclades within the section (Bruyns et al. 2017). Many non-trap flowered species in sect. Chamaesiphon convergently evolved morphological features considered typical for brood-site mimicking plants, i.e., prostrate or sub-erect stems, foul smelling, reddish-purplish or maroon speckled or striated flowers presented close to the ground (Johnson & Schiestl 2016; Fig. 1 A – L). Among the brood-site mimicking species in sect. Chamaesiphon, C. gilboaensis appears to be most closely related to C. ngomensis (Fig. 1 I)—a KwaZulu-Natal endemic species only known from the a rea of Ngome, about 220 km north-east of the Karkloof where C. gilboaensis occurs. Ceropegia ngomensis is the only other South African species presenting similarly structured steep cylindrical-campanulate corolla tubes in combination with prostrate stems. Preliminary unpublished molecular findings support a close relationship between both species (Heiduk et al. unpublished). Although C. gilboaensis also shares particular vegetative and floral characters with other brood-site mimicking members of sect. Chamaesiphon occurring in KwaZulu-Natal and adjacent regions, it may be distinguished as explicated below (see also Table 1). The prostrate stems with elliptic leaves of C. gilboaensis are distinct from the sub-erect to erect stems with commonly elliptic- or ovate-lanceolate leaves of C. alpina (Dyer 1977a: 18) Bruyns in Bruyns et al. (2017: 433), C. australis, C. campanuliformis, C. chlorozona, C. modestantha, C. perdita (Dyer 1977a: 10) Bruyns in Bruyns et al. (2017: 434), C. petrophila and C. spathulata (Lindley 1827: t. 1113) Bruyns in Bruyns et al. (2017: 435). The corolla of Ceropegia gilboaensis is, with rare exception (one plant found with sparse white trichomes on corolla lobe margins), glabrous, allowing for separation from C. alpina, C. australis, C. campanuliformis, C. chlorozona, C. perdita, C. petrophila, C. remota, C. swazica (Dyer 1976: 56) Bruyns in Bruyns et al. (2017: 435) and C. spathulata which all have distinctively puberulous, hirsute or pilose corolla lobes and/or vibratile hairs fringing the corolla lobes (Fig. 1, Table 1; Dyer 1980). In Ceropegia gilboaensis the corolla is steep cylindrical-campanulate (Figs 3 A – B, F, 4 A – D), which is distinct from the bowl-shaped corolla of C. chlorozona (Fig. 1 F) and the flat or shallowly saucer-shaped corollas of C. bruceae (Fig. 1 E), C. coddii (Fig. 1 G), and C. pulchellior (Fig. 1 K). Campanulate corollas are also characteristic of C. alpina, C. australis (Figs 1 A – D), C. modestantha (Fig. 1 H), C. perdita, C. petrophila (Fig. 1 J), C. remota (Fig. 1 L) and C. spathulata. However, in these species the corollas are less steep and the corolla lobes generally proportionally longer than the fused portion of the corolla (ca. ±1/ 2 in C. gilboaensis, see Table 1); in C. campanuliformis, the corolla lobes are proportionally shorter (±1/3) than the fused part of the corolla. The follicles of Ceropegia gilboaensis are verrucose, while they are smooth in all other species discussed here. Generally, raucous, non-smooth follicles are rather rare in Ceropegia although known for example from the succulentleaved climbers C. sandersonii Hooker (1869: t. 5792), which has follicles markedly verrucose to tuberculate, and C. cimiciodora Obermeyer (1933: t. 488) in which species they are thinly tuberculate (Dyer 1980). Within sect. Chamaesiphon, verrucose follicles have not earlier been reported. Ceropegia pulchellior and C. modestantha are two KwaZulu-Natal endemics that grow in rocky grasslands in relatively close proximity to C. gilboaensis (Fig. 2), which in view of their superficial similarity could lead to confusion with C. gilboaensis. However, in addition to the differences mentioned above, C. gilboaensis can be distinguished from these species as follows: when viewed from above, the sepals of C. pulchellior protrude from beneath the fused corolla portion, a feature not seen in C. gilboaensis. The spreading corolla lobe tips are basally ovate and distally attenuate-tapering and often reflexed in C. modestantha but are clearly triangular in C. gilboaensis. In C. pulchellior the gynostegium projects from the centre of the corolla whereas in C. gilboaensis, as also in C. modestantha, the gynostegium is sunk into the fused section of the corolla. The margin formed by the outer corona lobes is U-shaped in both C. modestantha (Fig. 5 C) and C. pulchellior (Fig. 5 D), while it is V-shaped in C. gilboaensis (Fig. 5 A). The outer corona lobes are lower than the style-head in C. modestantha (Fig. 5 C) and C. pulchellior (Fig. 5 D) but are level with the style-head in C. gilboaensis (Fig. 5 A). Both Ceropegia modestantha and C. pulchellior occur on Natal-Group sandstone while C. gilboaensis occurs on dolerite, as does C. ngomensis. Ceropegia modestantha is relatively widely distributed, ranging from near Nkandla in the north, to the Little Noodsberg, and south to the Umzinto coastal escarpment (Dyer 1980; Styles & Wragg 2008). Ceropegia pulchellior has a more limited distribution than C. modestantha, occurring between Pietermaritzburg and Durban along a narrow line of suitable habitat (Styles & Wragg 2008), with an outlier subpopulation to the south, “on an isolated mountain top near Eston…” (Styles & Wragg 2008: 46), within the range of C. modestantha. There, both species apparently co-occur and probably hybridize (Styles & Wragg 2008). A third species occurring relatively close to C. gilboaensis, but ranging somewhat further to the northeast is C. chlorozona, which grows on dolerite as well as sandstone. Other species in sect. Chamaesiphon share vegetative and/or floral characteristics with C. gilboaensis but are geographically separated from it. From north to south these species occur as follows: C. bruceae and C. coddii on the Mpumalanga escarpment and in areas of north-eastern eSwatini (Dyer 1980), with C. coddii also being recorded in KwaZulu-Natal (pers. obs.); C. swazica in eSwatini and adjacent Mpumalanga; C. remota in the Utrecht-Wakkerstroom-Paulpietersburg areas of KwaZulu-Natal; C. petrophila at Boston, Byrne and at Underberg in KwaZulu-Natal; C. perdita in the Drakensberg (Dyer 1980) above 2000 m near the trijunction of KwaZulu-Natal, Eastern Cape and Lesotho; C. alpina above 2300 m in southern Lesotho; C. campanuliformis and C. spathulata in the Eastern Cape; C. australis at Port Shepstone and as far south as Port St. Johns, restricted to Msikaba Formation sandstone. This lastmentioned species shows remarkable variation in flower colour and the length of the corolla lobes (Figs 1 A – D; see also Dyer 1983), even within the same population (Figs 1 A – C)., Published as part of Heiduk, Annemarie, Crouch, Neil R. & Styles, David G. A., 2023, Ceropegia gilboaensis (Apocynaceae), a new species from the Midlands of KwaZulu-Natal, South Africa, pp. 125-136 in Phytotaxa 591 (2) on pages 129-134, DOI: 10.11646/phytotaxa.591.2.4, {"references":["Trinius, C. B. & Ivanovich, F. J. (1842) Aristida junciformis Trin. & Rupr. Species Graminum Stipaceorum 143.","Steudel, E. G. (1855) Synopsis Plantarum Glumacearum. Pars I. Gramineae. J. B. Metzler, Stuttgartiae, 475 pp.","Durand, T. A. & Schinz, H. (1894) Sporobolus natalensis T. Durand & Schinz. Conspectus Florae Africae 5: 822.","Klopper, R. R. & Smith, G. F. (2010) Aloe neilcrouchii, a new robust Leptaloe from KwaZulu-Natal, South Africa. Bothalia 40 (1): 95 (- 96; fig. 34, map).","Bolus, H. (1881) Crassula dependens Bolus. Journal of the Linnean Society, Botany 18: 391.","Baker, J. G. (1870) Monograph of Scilla: sections Ledebouria and Drimiopsis. Saunders Refugium Botanicum 3: App. 5.","Venter, S. & Edwards, T. J. (2003) Ledebouria sandersonii (Baker) S. Venter & T. J. Edwards. Bothalia 33 (1): 50.","South African National Biodiversity Institute (2006 - 2018) The Vegetation Map of South Africa, Lesotho and Swaziland. Mucina, L., Rutherford, M. C. & Powrie, L. W. (eds.) Version 2018. [online] Available from: http: // bgis. sanbi. org / Projects / Detail / 186 (Accessed 31 March 2023)","Forsskal, P. (1775) Themeda triandra Forssk. Flora Aegyptiaco-Arabica 178.","Mucina, L., Hoare, D. B., Lotter, M. C., Du Preez, J., Rutherford, M. C., Scott-Shaw, C. R., Bredenkamp, G. J., Powrie, L. W., Scott, L., Camp, K. G. T., Cilliers, S. S., Bezuidenhout, H., Mostert, T. H., Siebert, S. J., Winter, P. J. D., Burrows, J. E., Dobson, L., Ward, R. A., Stalmans, M., Oliver, E. G. H., Siebert, F., Schmidt, E., Kobisi, K. & Kose, L. (2006) Grassland Biome. In: Mucina, L. & Rutherford, M. C. (eds.) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia, Pretoria, South Africa, pp. 349 - 436.","Van Wyk, A. E. & Smith, G. F. (2001) Regions of floristic endemism in southern Africa. A review with emphasis on succulents. Pretoria: Umdaus Press, 199 pp.","IUCN Standards and Petitions Committee (2019) Guidelines for Using the IUCN Red List Categories and Criteria. Version 14. Prepared by the Standards and Petitions Committee, 113 pp. Available from: http: // www. iucnredlist. org / documents / RedListGuidelines. pdf (accessed 22 September 2020).","Bruyns, P., Klak, C. & Hanacek, P. (2017) A revised, phylogenetically-based concept of Ceropegia (Apocynaceae). South African Journal of Botany 112: 399 - 436. https: // doi. org / 10.1016 / j. sajb. 2017.06.021","Johnson, S. D. & Schiestl, F. P. (2016) Floral Mimicry. Oxford University Press, 256 pp.","Dyer, R. A. (1977 a) New records of Brachystelma. Journal of South African Botany 34 (1): 9 - 20.","Lindley, J. (1827) Brachystelma spathulatum Lindl. Botanical Register 13: t. 1113.","Dyer, R. A. (1976) Asclepiadaceae. Bothalia 12 (1): 53 - 64.","Dyer, R. A. (1980) Asclepiadaceae (Brachystelma, Ceropegia, Riocreuxia). In: Leistner, O. A. (ed.) Flora of Southern Africa 27 (4). Botanical Research Inst., Pretoria, pp. 1 - 91.","Hooker, J. D. (1869) Ceropegia sandersonii - Mr. Sanderson´s Ceropegia. Botanical Magazine 95: t. 5792.","Obermeyer, A. A. (1933) Ceropegia cimiciodora Oberm. Flowering Plants of South Africa xiii. t. 488.","Styles, D. G. & Wragg, P. D. (2008) Notes on Brachystelma pulchellum and B. modestum. Asklepios 100: 42 - 53.","Dyer, R. A. (1983) Ceropegia, Brachystelma and Riocreuxia in Southern Africa. Balkema, Rotterdam, 242 pp."]}

Identification key for males of southern African sisyphine species 1. Lateral ridge on the pronotal disc extends from the posterior margin without reaching the lateral margin (Fig. 1F); strong setae absent on the external margin of the first meso- and metatarsal segments (Fig. 1D)........................ Neosisyphus (2) ��� Lateral prothoracic ridge between pronotal disc and prothoracic episternum complete (Fig. 1E); first tarsal segments of the meso- and meta-thoracic legs with a row of dense setae in a straight line on the external edge (Fig. 1C)........................................................................................................................... Sisyphus (13) 2. Mesotibia strongly modified by indentations and protrusions; a large spine distally on the posterior margin of the mesofemur. Completely black, pygidium simple and lacking a pair of distinct tubercles. Eastern coastal forests of South Africa and South Mozambique................................................................................................................(tibialis species-group; Neosisyphus mirabilis (Arrow,1927) (Figs. 23, 24A) ��� Mesotibia simple, without any spine projected distally on the posterior margin of the mesofemur.................................................................................................(3) 3. Femur and tibia of mesolegs without cuticular projections on the posterior margin............................................. barbarossa species-group (Figs. 17 A���D) (4) ��� Femur and tibia of mesolegs with cuticular projections on the posterior margin...(7) 4. Metafemur with sharp pointed spine on the mid-posterior margin (Figs. 17 A���B)........................................................................................................................(5) ��� Metafemur with angled-protuberance on the mid-posterior margin (Figs. 17 C���D)........................................................................................................................(6) 5. Projecting metatrochanter longer than �� of the metafemur length in major males and �� of the metafemur length in minor males. In major males the metatrochant- er is slightly inwardly-curved at the tip and straight in minor males; setae on the elytra black, recurved at the apex. Lowland dry savannas in southern Africa.................................................. Neosisyphus calcaratus (Klug, 1855) (Figs. 17, 18B) ��� Projecting metatrochanter shorter in minor males ( Neosisyphus barbarossa (Wiedemann, 1823) (Figs. 17, 18A) 6. Projecting metatrochanter relatively short; four times shorter than total length of metafemur. Lowland and upland moist vegetation from northeast to southern Africa..................................... Neosisyphus setiger (Roth, 1851) (Figs. 17, 18D) ��� Metatrochanter slighty inwardly-projected at the apex, metatrochanter projection is �� the length of the metafemur. Upland woodland in southern Angola............................................. Neosisyphus tembyi Daniel & Davis sp. n (Figs. 17, 18C) 7. Posterior margin of the mesoleg with spine or projection distally on mesofemur and basally on mesotibia; first quarter of mesotibia from femoro-tibial joint abruptly narrowed................................ spinipes species-group (Figs. 20 A���C) (8) ��� Mesotibial spine projecting posteriorly and gradually narrowing distally........(10) 8. Spine present on the mid-posterior margin of the metafemur; metatrochanter relatively short, �� of the metafemur length........................................................... (9) ��� Spine absent on posterior margin of the metafemur; metatrochanter is �� the length of the metafemur in major males. Shaded vegetation and thickets from east to southern Africa........ Neosisyphus fortuitus (P��ringuey, 1901) (Figs. 20, 21A) 9. Distinct triangular shaped projection present on the basal posterior margin of the mesotibia. Parameres not curved at the apex (Fig. 21C). Moist upland and coastal grassland from east to southern Africa............................................................................................................. Neosisyphus spinipes (Thunberg, 1818) (Fig. 20C) ��� Lacking a triangular protuberance on the basal posterior margin of the mesotibia in minor males. There is a slight sinuosity in major males. Parameres curved downward apically (Fig. 21B). Lowland dry savanna from east to southern Africa................................................. Neosisyphus infuscatus (Klug, 1855) (Fig. 20B) 10. Basal posterior spine on mesotibia projecting perpendicularly; body uniformly brown; size of beetles 6.0��� 10 mm......... rubrus species-group (Figs. 29 A���B) (11) ��� Basal posterior spine on mesotibia not projecting perpendicularly to the ventral apical margin of the femur; colour is not uniform overall, size of beetles 11.0��� 13.5 mm...................................... quadricollis species-group (Figs. 25 A���B) (12) 11. Parameres with protrusion apical-laterally (Fig. 30B). Upland and coastal moist grassland and savanna in southern Africa......................................................................................................... Neosisyphus rubrus (Paschalidis, 1974) (Fig. 29B) ��� Parameres simple (Fig. 30A). Cool, dry savanna, karoo and grassland in South Africa (Northern Cape and Free State) and Namibia (Etosha Pan, Okaukuejo)..................................... Neosisyphus macrorubrus (Paschalidis, 1974) (Fig. 29A) 12. Lateral margin of elytra emarginate; concavity of margin distinctly deep in the middle; short projecting metatrochanter in males (1/3 of metafemur length); metafemur lacking spines. Deep sands and shrubland of southwest coast, and karoo in the Eastern Cape (South Africa)................................................................................................... Neosisyphus quadricollis (Gory, 1833) (Figs. 26, 27A) ��� Margin of elytra with regular shape; a protuberance distally on the posterior margin of the metafemur, the metatrochanter exceeds �� the length of the metafemur in males. Moist highland grassland in South Africa............................................................................................. Neosisyphus kuehni (Haaf, 1955) (Figs. 26, 27B) 13. Setae on the elytral interstriae arranged in tufts............................................................................................................. muricatus species-group (Figs. 2 A���F) (14) ��� Setae on the elytral interstriae evenly distributed, not grouped in tufts.......... (20) 14. Setae arranged in tufts on the pronotum, which alternate with depressed bare patches.......................................................................................................... (15) ��� Setae on the pronotum not arranged in tufts................................................. (17) 15. Concave and upwardly curved margin between the anterior medial teeth of the clypeus; macropterous (Fig. 2I). Woodland thickets and forest in north South Africa and Swaziland.......... Sisyphus fasciculatus Boheman, 1857 (Figs. 2, 3C) ��� Anterior margin between medial clypeal teeth not upcurved; hind wings reduced or completely brachypterous (Figs. 2 H���J).....................................................(16) 16. Anterior margin between medial clypeal teeth concave; pronotum wider anteriorly than posteriorly. Parameres (Fig. 3F). Only known from Swaziland........................................................................ Sisyphus swazi Daniel & Davis sp. n (Fig. 2F) ��� Straight edge between medial teeth of clypeus; width of pronotum uniform from anterior to posterior margins. Parameres (Fig. 3G). Popa Falls forest (Namibia), Central Mozambique and East Africa.................................................................................................................... Sisyphus alveatus Boucomont, 1935 (Fig. 2G) 17. Very strong lateral emargination and distinct shiny punctation on pronotum. Cooler southern forest and montane grassland in South Africa............................................................................ Sisyphus muricatus (Olivier, 1789) (Figs. 2, 3A) ��� No distinct lateral emargination on the pronotum.........................................(18) 18. Punctation on the pronotum unclear; interstriae 1, 3, 5, and 7 with rows of black setae arranged in tufts, interstriae 2, 4, and 6 with sparsely arranged single setae. Coastal sand forest of northeast South Africa and southeast Mozambique................................ Sisyphus neobornemisszanus Daniel & Davis, 2016 (Figs. 2, 3B) ��� Punctation on the pronotum conspicuous; setae on pronotal disc arise from the centre of ocellate punctures (Fig. 5H)............................................................(19) 19. Proximal elytral setae evenly arranged; mid-basal elytral setae arranged in tufts of scattered individual setae. Pronotal disc with three prominent round depressions. Parameres (Fig. 3E). Primarily upland and highland grassland in Gauteng, Kwa- Zulu Natal, Limpopo, Mpumalanga and Northwest Province..................................................................................... Sisyphus manni Montreuil, 2015 (Fig. 2E) ��� Elytra entirely covered in tufts of setae, each tuft dense and compact; depressions on the pronotal disc absent; body larger than S. manni and oval. Parameres (Fig. 3D). Unshaded vegetation in the Eastern Cape......................................................................................... Sisyphus perissinottoi Montreuil, 2015 (Fig. 2D) 20. Well-developed tufts of epipleural setae (Fig. 5H); edge between medial teeth almost straight; setae on the pronotum evenly distributed........................................................................................................ goryi species-group (Figs. 5 A���E) (22) ��� Weak tufts of epipleural setae; margin between medial teeth V-shaped)................................................................................................. umbraphilus species-group 21. Pronotal disc with setae interspersed with bald patches. Dense woodland and thicket in KwaZulu Natal, Gauteng and North West Provinces (South Africa)......................................... Sisyphus umbraphilus Daniel & Davis, 2016 (Fig. 8A) ��� Pronotal disc without clear setae interspersed with bald patches.................... (26) 22. Setae on the pronotum inserted between or on the posterior margin of ocellate punctures (Fig. 5F)........................................................................................ (23) ��� Setae on the pronotum inserted centrally on ocellate punctures.................................................................................................................................. (Fig. 5G) (25) 23. Fine setae on elytra arranged in single rows, epipleural tufts containing few setae ( Sisyphus caffer Boheman, 1857 (Figs. 5, 6B) ��� Much finer setae on elytra, distributed in single rows; epipleural tufts containing many setae (>10); elongate body shape..........................................................(24) 24. Setae on the pronotum and elytra of uniform colour and regularly arranged. Parameres (Fig. 6D). Widespread woodland species in sub-Saharan Africa................................................................................ Sisyphus goryi Harold, 1859 (Fig. 5D) ��� Setae on pronotum and elytra bicoloured (black and gray). Parameres (Fig. 6A). Open woodland in southern Angola........................................................................................................... Sisyphus bicuariensis Daniel & Davis, sp. n (Fig. 5A) 25. Elytral setae thick, arranged in non-single rows and forming bunches; body size: 6.6��� 4.4 mm. Open lowland vegetation in South Africa, Mozambique, and Zimbabwe....................................... Sisyphus sordidus Boheman, 1857 (Figs. 5, 6E) ��� Elytral setae fine, regularly arranged on interstriae; body size: 10���8.0 mm. East African miombo woodland; known only from Quirimbas National Park (Northern Mozambique)..................... Sisyphus genierorum Montreuil, 2015 (Figs. 5, 6C) 26. Setae on outer margin of elytra not arranged in tufts; margin between medial teeth concave or straight; very large ocellate punctures on dorsal surface of pronotal disc...................................................... costatus species-group (Figs. 11 A���E) (27) ��� Setae on outer margin of elytra not arranged in tufts; margin between medial teeth narrowed (almost V-shaped); very fine ocellate punctures on dorsal surface of pronotal disc......................................... seminulum species-group (Figs. 6 A���C) (30) 27. Dorsal surface of the clypeus and frons with inconspicuous ocellate punctation, but more visible on the vertex (Figs. 11 G���I).................................................. (28) ��� Dorsal surface of the head (clypeus, frons and vertex) with distinct ocellate punctation (Figure11J).......................................................................................... (30) 28. Vertex with dense ocellate punctation, margin between medial teeth straight; genae convex anteriorly (Fig. 11I); elytral setae fine and uniformly arranged. Moist savanna or dense woodland and riverine vegetation in the interior of South Africa. Shaded vegetation on coastline from Sofala Bay (Mozambique) to Eastern Cape (South Africa)................ Sisyphus inconspicuus Daniel & Davis, sp. n (Fig. 5B) ��� Vertex with scattered ocellate punctation (Fig. 11 G���H), elytral setae thicker, alternating on the interstriae between rows of well-developed and less developed setae (Fig. 11F)......................................................................................................(29) 29. Margin between medial teeth distinctly concave and upcurved; lacking carinae on the frons; genae virtually straight laterally (Fig. 11G). Parameres (Fig. 12B). Highland grassland in South Africa....................................................................................................................... Sisyphus costatus (Thunberg, 1818) (Fig. 11B) ��� Edge between medial teeth straight; well-defined clypeo-frontal carina; genal margin virtually convex (Fig. 11H). Parameres (Fig. 12E). Shaded vegetation from Western and Eastern Cape Provinces, South Africa.......................................................................................... Sisyphus australis Daniel & Davis sp. n (Fig. 11E) 30. Small-bodied; genae with shallow depression dorsally; pronotal disc showing distinct pattern of bare patches amongst sparse short setae; metafemur simple. Highland forest under cooler conditions in Zimbabwe and Malawi........................................................................ Sisyphus gazanus Arrow, 1907 (Figs. 11, 12C) ��� Large-bodied; genal depression absent; setae arranged regularly on pronotal disc; anvil- shaped projection on mid-posterior margin of metafemur. Lowland shaded Savanna from east to southern Africa..................................................................................................... Sisyphus impressipennis Lansberge, 1886 (Figs. 11, 12A) 31. Relatively large-bodied; pronotal disc with conspicuous metallic sheen; open woodland in north of Namibia and south of Angola............................................................................................ Sisyphus splendidus Montreuil, 2015 (Fig. 14A) ��� Relatively small to medium-bodied................................................................ (32) 32. Very small-sized body (3.0��� 3.8 mm); pronotal disc dull bearing some depressions. Shaded vegetation in eastern and southern Africa....................................................................................... Sisyphus nanniscus P��ringuey, 1901 (Figs. 14C, 15B) ��� Medium-sized body (3.9���5.0 mm)................................................................ (33) 33. Pronotal disc bears sparse, long and fine yellow setae, separated by linear bare patches (Fig. 14E); parameres notched dorsally (Fig. 15A). Dense coastal woodlands and forest from northeast South Africa to southern Mozambique........................................................ Sisyphus oralensis Daniel & Davis, 2016 (Fig. 14B) ��� Pronotum bears thick and short golden setae; directional projection of the setae well-defined (Fig. 14F); parameres without excavation dorsally, attenuating sharply at the apex (Fig. 15C). Forest and dense woodland species in central Mozambique................... Sisyphus auricomus Daniel & Davis sp. n (Fig. 14D), Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 5-9, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Klug, J. C. F. (1855) Diagnosen neuer Coleoptera aus Mossambique. Bericht uber die zur Bekanntmachung geeigneten Verhandlungen der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 643 - 660.","Wiedemann, C. K. W. (1823) Zweihundert neue Kafer von Java, Bengalen und dem Vorgebirge der guten Hoffnung. Zoologisches Magazin, 2 (1): 3 - 133.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Thunberg, C. P. (1818) Coleoptera Capensia, Antennis Lamellatis, sive, clava fissili instructa, descripta, memoires de l'Acad. T. VI.","Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp.","Gory, M. (1833) Monographie du genre Sisyphe. Mequignon-Marvis pere et fils, Paris. 1 - 15 p.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Boheman, C. H. (1857) Insecta Caffrariae annis 1838 - 1845 a JA Wahlberg collecta. Pars. II. Coleoptera (Scarabaeides). Holmiae: Offieina Nordstedtiana. - 395 pp, 1.","Boucomont, A. (1935) Coleoptera. VI. Scarabaeidae Scarabaeini. In: Jeannel, R. (Ed.), Mission Scientifique de l'Omo. Tome 2 Zoologie, Fascicule 16. Paul Chevalier & Fils, Paris, France, pp. 279 - 290.","Olivier, M. (1789) Entomologie, Histoire Naturelle des Insectes, Avec leurs caracteres generiques et specifiques, leur description, leur synonymie, et leur figure enluminee. Coleopteres, Tome premier, Paris, 1 - 455 pp.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Harold, E. von (1859) Beitrage zur Kenntniss einiger coprophagen Lamellicornien. Deutsche Entomologische Zeitschrift, 3 (2 - 3), 193 - 224.","Lansberge, J. W. (1886) Scarabaeides, Buprestides et Cerambycides de l'afrique occidentale. Notes from Leyden Museum, 8 (2): 69 - 120."]}

3. Gaylussacia Kunth (1818: 215). Type:— Gaylussacia buxifolia Kunth (1818: 276). Subshrubs to shrubs, erect to procumbent, orthocladous to deliquescent. Leaves alternate, papery to coriaceous; blade glabrous to tomentose; margin entire to serrate; apical callous glandular; venation camptodromous. Racemes axillary and subapical with bracts at the base of the inflorescence axis; floral bract 1, conspicuous at pedicel base; bracteoles 2 along the pedicel. Flowers 5-merous; calyx lobes basally connate and pubescent to glabrous; corolla gamopetalous, campanulate to urceolate; stamen 10, epipetalous; staminal filaments straight, flat, pubescent to tomentose; anthers dorsifixed; thecae parallel, long-tubular, dehiscence poricidal or by a small apical cleft; ovary inferior; pseudo-10- locular, one ovule per locule; stigma depressed-capitate. Fruit drupoid (nuculanium), globose; seeds lenticular. The genus Gaylussacia in Rio Grande do Sul comprises three species distributed mainly in a highland grassland of Campos de Cima da Serra (Figs. 23 A, B, C), also reaching the regions of Depressão Central (Fig. 23 A) and South Coast (Fig. 23 B). Gaylussacia are associated with forest edges and in grasslands along cloud forests and riverine forests (Fig. 17 B; 19 B, E; 21 A), in clay and acidic soils (Fig. 17 A), temporary wetlands and peat bogs (Fig. 19 A) in rocky hillsides and cliffs, basaltic outcrops, wet sandy soils, peat bogs and sandy soils at restinga., Published as part of Dalastra, Claudenice Hilda & Heiden, Gustavo, 2022, Ericaceae in Rio Grande do Sul state, Southern Brazil, pp. 149-190 in Phytotaxa 564 (2) on page 174, DOI: 10.11646/phytotaxa.564.2.2, http://zenodo.org/record/7087201, {"references":["Kunth, C. S. (1818) Ericaceae. In: von Humboldt, F. H. A., Bonpland, A. J. & Kunth (eds.) Voyage de Humboldt et Bonpland. Nova Genera et Species Plantarum 3: 215 - 250."]}

6. Sida farroupilhensis Krapovickas & Bueno (2014: 87) (Figures 4A–F, 2, 14C). Holotype:— BRAZIL. Rio Grande do Sul: Farroupilha, “Parque dos Pinheiros, Parque dos Pinheiros, erva com aproximadamente 1 m de altura, bastante frequente nos lugares úmidos e sombreados, flores amarelas que se fecham ao ser coletadas”, 23 May 1978, O. Bueno 676 (HAS!; isotype CTES!). Taxonomic notes: — Sida farroupilhensis is distincted from other species of section Sidae (Table 1) by the presence of aristae smaller than the mericarp body, mericarps dehiscent only at the apex, solitary flowers without bifid bracts, leaf blades ovate-rhombic and with simple trichomes in adaxial surface (Figures 4A–F, 14C) (Krapovickas 2014). A relevant diagnostic feature of the species is the reticulate leaf’s abaxial surface, formed by the tertiary veins, which is covered by sparse, small stellate trichomes. Calyx also with tertiary veins forming a characteristic reticulate and with very sparse small stellate trichomes. In the present work, photos of the species in its habitat are presented for the first time. Distribution, habitat, and conservation status: —The species is endemic to Araucaria Forest of northeast of Rio Grande do Sul, Brazil (Figure 2), in the edges or understory of Forests, and rarely in Highland grassland. With an EOO of 3,813.842 Km² and an AOO of 40 km ² S. farroupilhensis could be considered Endangered (EN). However, the species is usually abundant in places where it occurs and does not show subpopulations, locations, EOO and AOO declines or fluctuations. That is, the species does not meet the conditions B (a), B(b) and B(c) and is considered Least Concern (LC). Representative specimens: — BRAZIL. Rio Grande do Sul. Canela, Flona, WGS 84 - 29.321389°S - 50.816667°W, 23 March 2019, F . Gonzatti 5061 (HUCS); Caxias do Sul, Bairro Nossa Senhora do Rosário, WGS 84 - 29.130833°S - 51.154167°W, 29 September 2019, M . Grings 2078 (ICN); Gramado, Bairro Moura, 06 May 2020, M . Grings 2130 (ICN); São Francisco de Paula, próximo ao lago São Bernardo, WGS 84 - 29.459764°S - 50.570602°W, 08 June 2012, M . Grings 2112 (ICN); Vacaria, Parque de Rodeios, WGS 84 - 28.481828°S - 50.910969°W, 27 April 2019, M . Grings & A. S. Mello 2034 (ICN)., Published as part of Grings, Martin & Boldrini, Ilsi Iob, 2022, Synopsis of Sida (Malvoideae, Malvaceae) in the state of Rio Grande do Sul, Brazil, pp. 105-135 in Phytotaxa 542 (2) on page 114, DOI: 10.11646/phytotaxa.542.2.1, http://zenodo.org/record/6415487, {"references":["Krapovickas, A. (2014) Nuevas especies de Sida Seccion Sida (Malvaceae). Bonplandia 23: 65 - 118. https: // doi. org / 10.30972 / bon. 232258"]}

21. Sida tuberculata R.E. Fries (1908: 33) var. pseudo-rhombifolia Monteiro (1949: 522) (Figures 8, 11L–Q, 14T) Lectotype (designated by Krapovickas 2014: 111):— BRAZIL. Minas Gerais: Belo Horizonte, 29-I-1934. H. Monteiro 711 (RBR!, isotype US!). Taxonomic notes: — S. tuberculata var. pseudo-rhombifolia belongs to section Sidae (Table 1) and is a species morphologically related to Sida viarum and Sida rhombifolia (Krapovickas 2014, Monteiro-Filho 1949). It differs from these species by the presence of 6–8 mericarps (rarely nine), vs. 5 mericarps in S. viarum and 9–14 mericarps in S. rhombifolia; presence of simple trichomes on the margins of the calyx lobes; flowers solitary or crowded in subsessile axillary glomerules, the pedicels up to 15 mm long on solitary flowers (Figures 11L–Q, 14T) (Krapovickas 2005, Brandão et al. 2017). Distribution, habitat and conservation status: —The species occurs in Venezuela, Bolivia, Paraguay, Argentina, Uruguay and Brazil (northeast, midwest, southeast and south) (Krapovickas 2014, Bovini 2020). In Rio Grande do Sul the species occurs in Park grassland, Sandy grassland, Shallow soil grassland, Inland sub-montane grassland, Bush grassland, Highland grassland, Aristida grassland, Coastal grassland and Inland subtropical Forest (Figure 8). The species is here considered Least Concern (LC) because it has large EOO and AOO, since it is a frequent and ruderal species, and widely distributed. Representative specimens: — BRAZIL. Rio Grande do Sul. Alegrete, Estação do Tigre, 23 November 1958, J . Mattos 6223 (HAS); Barra do Quaraí, Parque Estadual do Espinilho, 54 m, WGS 84 S -30.202154° W - 57.505048°, 02 December 2020, M . Grings et al. 2197 (ICN); Caçapava do Sul, defronte a Pedra do Segredo, 25 March, 1995, O . Bueno et al. 4198 (HAS); Caxias do Sul, Água Azul, 10 January 1981, O . Bueno 2882 (HAS); Giruá, Granja Sodal, 04 April 1966, K . Hagelund 4223 (HAS); Porto Xavier, 28 Km SE de Porto Xavier, 18 February 1978, A . Krapovickas & C. L. Cristóbal s.n. (ICN 45915); Cristal, 7 Km da ponte de divisa com Camaquã, BR-116, 07 January 1980, O . Bueno 2064 (HAS); São Pedro do Sul, 03 December 1981, O . Bueno 3425 (HAS)., Published as part of Grings, Martin & Boldrini, Ilsi Iob, 2022, Synopsis of Sida (Malvoideae, Malvaceae) in the state of Rio Grande do Sul, Brazil, pp. 105-135 in Phytotaxa 542 (2) on page 130, DOI: 10.11646/phytotaxa.542.2.1, http://zenodo.org/record/6415487, {"references":["Fries, R. E. (1908) Studien uber die amerikanische Columniferenflora. Kungliga Svenska Vetenskapsakademiens Handlingar 42: 1 - 67 + 7 plates.","Krapovickas, A. (2014) Nuevas especies de Sida Seccion Sida (Malvaceae). Bonplandia 23: 65 - 118. https: // doi. org / 10.30972 / bon. 232258","Monteiro-Filho, H. da C. (1949) As especies argentinas, brasileiras e uruguayas da seccao Malvinda do genero Sida (Malvaceas). Lilloa 17: 501 - 522.","Krapovickas, A. (2005) Malvaceae. In: Bacigalupo, N. M. (Ed.) Flora Ilustrada de Entre-Rios. 6 (4 b). Colecc. Ci. Inst. Nac. Tecnol. Agropecu, pp. 292 - 333.","Brandao, J. L., Baracho, G. S., De Sales, M. F. & Filho, M. P. V. (2017) Synopsis of Sida (Malvaceae, Malvoideae, Malveae) in the state of Pernambuco, Brazil. Phytotaxa 307 (3): 205 - 227. https: // doi. org / 10.11646 / phytotaxa. 307.3.5","Bovini, M. G. (2020) Sida in Lista de Especies da Flora do Brasil. Jardim Botanico do Rio de Janeiro. Available from: http: // floradobrasil. jbrj. gov. br / jabot / floradobrasil / FB 9206 (accessed 7 July 2020)"]}

13. Sida potentilloides A. Saint-Hilaire (1825: 178) (Figures 5, 7A–E, 14M). Lectotype (designated by Rodrigo 1944: t. 22):— BRAZIL. Rio Grande do Sul. “Secus silvam prope São Nicolao, in provincia Missionum”, s.d., A. St.-Hilaire C2/2708 bis (P barcode 02285462! [digital image]; isolectotypes P barcode 02285461! [digital image], P barcode 02285460! [digital image], MPU barcode 016972! [digital image], US barcode 00097962! [digital image]). = Sida anarthra Ekman (1910: 15) Lectotype (designated here):— ARGENTINA. Misiones. Posadas, Bonpland, in campo humido in marginibus silvulae, etc, 15 January 1908, E.L. Ekman 145 (S 14-46283! [digital image]; isolectotype S 14-46282! [digital image], LD barcode 1691486! [digital image], LP barcode 004304! [digital image], NY barcode 00222011! [digital image]). Nomenclatural notes: — Ekman (1910) designated the collection Ekman 145 as type from Sida anarthra. However, there are five specimens of this collection, all syntypes, deposited in herbaria S, LD, LP and NY. The specimen S 14-46283 was designated as a lectotype, because it is representative of the species. Taxonomic notes: —According to the description of sections of Sida from Fryxell (1985) we included the species in section Sidae by the leaves basally entire and distally dentate (dentate almost to the base in this species), base cuneate, sometimes more or less rhomboid, and number of mericarps 7–11 (Table 1). Striking features of this species are leaves dentate almost to the base, blades generally narrowly elliptic, narrowly oblong or oblong, abaxial surface tomentose, with margins coarsely serrate; corolla dark yellow, and mericarps with long aristae, greater than 1 mm long (Figures 7A–E, 14M). In the present work, photos of the species in its habitat are presented for the first time. Distribution, habitat and conservation status: — Sida potentilloides occurs in Paraguay, Argentina, Uruguay, and in southeast, and south of Brazil (Zuloaga et al. 2008, Bovini 2020). In state of Rio Grande do Sul the species grows generally in Araucaria Forest, sometimes associated with Highland grassland, but also in Inland subtropical Forest and Bush grassland (Figure 5), in edges of Forests, in ruderal environments and shrubland. The species is Least Concern (LC) because it has large EOO and AOO, since it is a frequent and often ruderal species. Representative specimens: — BRAZIL. Rio Grande do Sul. Bagé, Galpão de Pedra, 03 November 2010, M . Grings & P. J. Silva-Filho 1113 (ICN); Fontoura Xavier, 696 m, - 29.088611S - 51.029444W, 07 November 2018, M . Grings et al. 1950 (ICN); Jaquirana, Passo do S, 839 m, 29º05’32.6”S 50º21’51.6”W, 27 November 2009, M . Grings 819 (ICN); Nonoai, March 1945, B . Rambo s.n. (PACA 28083); Nova Petrópolis, Morro Korb, 700m, 02 November 2008, M . Grings 677 (ICN, HUCS); Tupanciretã, 26 January 1942, B . Rambo s.n. (PACA 9164)., Published as part of Grings, Martin & Boldrini, Ilsi Iob, 2022, Synopsis of Sida (Malvoideae, Malvaceae) in the state of Rio Grande do Sul, Brazil, pp. 105-135 in Phytotaxa 542 (2) on pages 119-120, DOI: 10.11646/phytotaxa.542.2.1, http://zenodo.org/record/6415487, {"references":["Saint-Hilaire, A. de, Jussieu, A. de & Cambessedes, J. (1825) Flora Brasiliae Meridionalis (quarto ed). A. Belin Bibliopolam, Parisiis, 395 pp.","Rodrigo, A. del P. (1944) Las especies argentinas y uruguayas del genero Sida (Malvaceae). Revista del Museo de La Plata (Nueva Serie), Seccion Botanica 6: 81 - 212.","Ekman, E. L. (1910) Beitrage zur Columniferenflora von Misiones. Arkiv for Botanik 9: 1 - 56.","Fryxell, P. A. (1985) The North and Central American species of Sida. Sida, Contributions to Botany 11: 62 - 91.","Zuloaga, F. O., Morrone, O. & Belgrano M. J. (2008) Catalogo de las plantas vasculares del Cono Sur (Argentina, sur de Brasil, Chile, Paraguay y Uruguay). Monographs in Systematic Botany from the Missouri Botanical Garden 107: 1 - 3348.","Bovini, M. G. (2020) Sida in Lista de Especies da Flora do Brasil. Jardim Botanico do Rio de Janeiro. Available from: http: // floradobrasil. jbrj. gov. br / jabot / floradobrasil / FB 9206 (accessed 7 July 2020)"]}

Austrolebias botocudo, new species (Figs. 1���2, Table 1) Holotype. MCP 54450, male, 47.8 mm SL, Brazil, Rio Grande do Sul, Vacaria, temporary pool near rio Socorro, rio Apua��-Inhandava drainage, upper rio Uruguay basin, 28��22���43.6��� S, 50��52���21.8��� W, altitude 934 meters a.s.l., L.E.K. Lan��s & M. V. Volcan, 26 July 2017. Paratypes. All from Brazil, Rio Grande do Sul, Vacaria, rio Socorro, rio Apua��-Inhandava drainage, upper rio Uruguay basin: MCP 54450, male, 51.6 mm SL, same locality as holotype, L.E.K. Lan��s, R.S. Godoy & V. Weber, 13 October 2016; MCP 54451, 2 males, 35.7���40.8 mm SL, collected with holotype; MCP 54452, 3 males, 44.7���55.6 mm SL, 3 females, 28.5���39.3 mm SL, same locality as holotype, L.E.K. Lan��s & M. V. Volcan, 26 Jul 2017; MCP 54453, 4 males, 36.5���54.6 mm SL (1 C&S), 6 females, 31.2���35.3 mm SL (1 C&S), 28��22���47.9��� S, 50��52��� 07.9��� W, altitude 926 meters a.s.l., L.E.K. Lan��s, R. S. Godoy & M. Foga��a, 23 November 2013; MCP 54454, 3 males, 37.4���43.5 mm SL, 1 female, 33.2 mm SL, 28��22���47.9��� S; 50��52���07.9��� W, altitude 926 meters a.s.l, L.E.K. Lan��s & R. S. Godoy & M. Foga��a, 30 Aug 2013; MCP 54455, 2 males, 28.0��� 34.7 mm SL (1 C&S), 1 female, 25.6 (C&S), 28��22���47.9��� S, 50��52���07.9��� W, altitude 926 meters a.s.l., L.E.K. Lan��s & A.B. Moraes, 10 May 2013; MCP 54456, 1 male, 34.8 mm SL (C&S), 1 female, 31.8 (C&S) 28�� 22���43.6��� S, 50��52���21.8��� W, altitude 934 meters a.s.l., L.E.K. Lan��s & A.B. Moraes, 10 May 2013. Diagnosis. Austrolebias botocudo differs from its congeners allocated in the subgenus Acrolebias by a unique male colour pattern consisting of body flank lilac grey with bluish white vertical bands (vs. flank pale yellowish brown, with dark brown bars in A. araucarianus; flank golden with purplish grey bars in A. carvalhoi; and flank dark bluish brown with yellowish golden or light grey vertical bands in A. nubium). The new species additionally differs from A. araucarianus and A. carvalhoi by males presenting melanophores irregularly distributed across the body (vs. melanophores absent in A. araucarianus and A. carvalhoi). Austrolebias botocudo differs from the remaining species of the subgenus Acrolebias, except from A. nubium, by presenting contact organs on anal fin in males (vs. contact organs absent). Austrolebias botocudo differ from A. nubium by the presence of discrete contact organs in the flank (vs. conspicuous contact organs), presence of generally only one contact organ by scale of lateral line (vs. two prominent contact organs by scale of lateral line), by presenting jaws slightly prognathous (vs. jaws short); by dorsal profile of head slight concave (vs. nearly convex), more neuromasts in the preopercular + mandibular series (36���43 vs. 26���35), a lower body depth in females (28.2���32.3 % SL vs. 32.6���37.8 % SL), and by presenting basihyal cartilage about 40���50% of basihyal length (vs. 60���70% of basihyal length). The new species differ from A. araucarianus by presenting neuromasts of the supraorbital series united (vs. first three neuromasts of the supraorbital series separated from the remaining neuromasts), preopercular and mandibular series of neuromasts united (vs. separated), by presenting pelvic fin well-developed (vs. pelvic-fin girdle rudimentary or absent), fewer scales in the longitudinal series (27���30 vs. 31���33), and by presenting more neuromasts in the ventral opercular series (2���4 vs. 1). Austrolebias botocudo differ from A. carvalhoi by presenting more neuromasts in the supraorbital (2���3+22���27 vs. 3+20) and lateral mandibular series (5���9 vs. 4), preopercular and mandibular series of neuromasts united (vs. separated), and by presenting basihyal cartilage about 40���50% of basihyal length (vs. 65% of basihyal length). Description. Morphometric data are presented in Table 1. Largest male examined 55.6 mm SL; largest female examined 39.3 mm SL. Dorsal profile slightly concave on head, convex from nape to the end of dorsal-fin base, nearly straight on caudal peduncle. Ventral profile convex from lower jaw to end of anal-fin base, about straight on caudal peduncle. Body moderately deep and compressed. Greatest body depth at vertical of pelvic fin origin. Lower jaw relatively long and prognathous, snout blunt. Dorsal-fin rays in males 22���24; in females 18���20. Dorsal fin sub-rectangular in males, with posterior tip rounded. Dorsal fin semi-circular in females. Origin of dorsal fin in males at vertical through neural spine of 7���9th vertebrae; in females through 10���11th vertebrae. Anal-fin rays in males 21���23, in females 18���20. Anal-fin sub-rectangular, with tip rounded in males. Anal-fin sub-triangular in females. Anteromedian rays of anal fin lengthened in females. Anal-fin origin at vertical through base of 3���5th dorsal-fin ray in males and through base of 2���4th dorsal-fin ray in females, and at vertical through pleural ribs of 8���9th vertebrae in males; and through 10���12th vertebrae in females. Caudal fin distal margin rounded, 24���27 rays in both sexes. Caudal-fin rays supported by last 4 vertebrae. Pectoral fin rays 12���14 in both sexes; pectoral-fin margin rounded. Pectoral-fin posterior tip reaching from before pelvic fin origin to 2nd anal-fin ray in males, from origin of pelvic fin or before it in females. Pelvic fins usually well-developed, with 5���6 rays, its tip extending from urogenital papilla to 2nd anal-fin ray in males, and to urogenital papilla in females. Urogenital papilla not attached to anal fin. Pelvic-fin bases medially separated by short interspace. Scales large and cycloid. Head and trunk entirely scaled, except ventral surface of head. Frontal squamation F, G, or H, sometimes irregularly arranged. No scales on dorsal and anal fin bases. Longitudinal series of scales 27���30, scales regularly arranged; transverse series of scales 11���16; 18���22 scale rows around caudal peduncle. Few, scattered contact organs on scales throughout body in males, when present located mainly at mid-ventral region of the flank from immediately distal tip of opercle to caudal peduncle. Usually one contact organ per scale of longitudinal series; rarely two contact organs along some scales of longitudinal series (present in only three males). Contact organs on distal portion of anteriormost 1���4 anal fin rays, present in 25% of males. Contact organs on outer region of 1���4 uppermost pectoral fin rays in males. Cephalic neuromasts: supraorbital 14���23, parietal 1���3, anterior rostral 1���2, posterior rostral 1, infraorbital 2���3+22���27, preorbital 2���3, otic 2���4, post-otic 2���5, supratemporal 1���2, median opercular 1, ventral opercular 2���4, preopercular + mandibular 36���43, lateral mandibular 5���9, paramandibular 1. Two neuromasts on caudal fin base. Six branchiostegal rays. Dermosphenotic ossification absent. Urohyal deep. Total number of vertebrae 28���29. Gill rakers in first branchial arch 3+8���9. Basihyal subtriangular, width about 60���75% of length; basihyal cartilage about 40���50% of total basihyal length. One to three teeth on second pharyngobranchial. Vomerine teeth absent. Colouration in life. Males (Fig. 2A���D). Ground colour of body lilac grey, brown in dorsal region, with 7���12 bluish white vertical bands, usually with narrower, overlapping or poorly defined stripes between well-defined bars. Some specimens in a dominant situation shown a darker coloured pattern, composed of a blue-purple background with vertical grey-green stripes. Pectoral and ventral region greyish. Melanophores irregularly distributed over the body, forming black dots, spots or irregular marks. Some specimens exhibit a concentration of melanophores in the middle of the flank, forming irregular black vertical bands. This pattern is more frequent in senile individuals. Opercle and preopercle greenish blue. Supraorbital and suborbital bars black. Pectoral fin hyaline, margin black. Pelvic fins dark blue or dark grey. Iris orange; dark vertical band crossing the eye. Dorsal fin dark blue with light grey elongated dots. Anal fin bluish-green with light, elongated grey dots; distal margin of anal fin darker. Caudal fin bluish grey with light grey dots and spots, distal margin hyaline. Females (Fig. 2E���H). Ground colour of body pale yellowish brown, generally with brown, dark brown or black spots distributed over the flank. Some individuals presenting a spotted pattern with several circular marks, which extends from dorsum to the medium flank. Dorsum generally pale yellowish brown. Some specimens present black melanophores in the dorsal region. Venter light yellowish grey. Sides of head yellowish brown, opercular region pale golden or bluish golden. Suborbital bar grey. Iris light yellow with transverse grey bar through centre of eye. Unpaired fins yellowish hyaline, with sparse and little conspicuous grey dots. Pectoral fin hyaline. Pelvic fin hyaline. Distribution. The species was recorded only in two temporary pools located in a highland grassland landscape at the rio Socorro, a tributary of upper rio Apua��-Inhandava (upper rio Uruguay basin), in altitudes between 926 and 934 meters a.s.l., Vacaria municipality, Rio Grande do Sul State, southern Brazil (Fig. 3). Etymology. The specific epithet ���botocudo��� is a reference to an indigenous people, which formerly inhabited the region and belonged to the Xokleng Lakl��n�� ethnic group, known during the 19th century as ���Botocudos���, because adult men used a botoque (a plug-like wooden disk) to enlarge the lower lip. An allusion to a feature presented by the species, where the lower jaw seems more prominent than other species belonging to the subgenus Acrolebias. Ecological notes. Austrolebias botocudo was found only in two isolated high-altitude temporary wetlands, located within a private farmland, with size areas of 2,500 and 13,000 m 2. These two biotopes present clear (sometimes turbid), still water, with a latosol soil type, and are located in a grassland landscape (Fig. 4). The wetlands are close to each other (less than one kilometer apart) but are hydrologically isolated. A portion of the larger temporary pool was excavated, probably to become a water source for cattle. There are small intermittent drainages associated with the pools. The vegetation is composed mainly by stands of Peruvian watergrass (Luziola peruviana), peatbog (Sphagnum sp.), and other emergent, floating and submerse macrophytes. During winter and early spring, when the temporary pools present water, the minimum air temperature reaches 0��C (often negative temperatures are recorded), and frost, dew, fog and even snow can occur. The area where the biotopes inhabited by A. botucudo lie within are used for extensive cattle ranching. In the surroundings, mainly soybean crops and orchards (apples and pears) have drastically altered the natural high-altitude grasslands landscape. Conservation. In accordance with IUCN criteria, Austrolebias botocudo should be considered as ���Critically Endangered���, with the criteria CR B2ab (ii, iii). The species presents an area of occupancy of 15,500 m 2 (AOO less than 10 km ��), present severely fragmented populations (a) with continued decline (b) in area of occupancy (ii) and quality of habitat (iii)., Published as part of Lan��s, Luis Esteban Krause, Volcan, Matheus Vieira & Maltchik, Leonardo, 2021, Two new annual fishes (Cyprinodontiformes: Rivulidae) unexpectedly discovered in the highlands of southern Brazil, pp. 499-520 in Zootaxa 4949 (3) on pages 501-506, DOI: 10.11646/zootaxa.4949.3.4, http://zenodo.org/record/4640565

Austrolebias botocudo, new species (Figs. 1–2, Table 1) Holotype. MCP 54450, male, 47.8 mm SL, Brazil, Rio Grande do Sul, Vacaria, temporary pool near rio Socorro, rio Apuaê-Inhandava drainage, upper rio Uruguay basin, 28°22’43.6” S, 50°52’21.8” W, altitude 934 meters a.s.l., L.E.K. Lanés & M. V. Volcan, 26 July 2017. Paratypes. All from Brazil, Rio Grande do Sul, Vacaria, rio Socorro, rio Apuaê-Inhandava drainage, upper rio Uruguay basin: MCP 54450, male, 51.6 mm SL, same locality as holotype, L.E.K. Lanés, R.S. Godoy & V. Weber, 13 October 2016; MCP 54451, 2 males, 35.7–40.8 mm SL, collected with holotype; MCP 54452, 3 males, 44.7–55.6 mm SL, 3 females, 28.5–39.3 mm SL, same locality as holotype, L.E.K. Lanés & M. V. Volcan, 26 Jul 2017; MCP 54453, 4 males, 36.5–54.6 mm SL (1 C&S), 6 females, 31.2–35.3 mm SL (1 C&S), 28°22’47.9” S, 50°52’ 07.9” W, altitude 926 meters a.s.l., L.E.K. Lanés, R. S. Godoy & M. Fogaça, 23 November 2013; MCP 54454, 3 males, 37.4–43.5 mm SL, 1 female, 33.2 mm SL, 28°22’47.9” S; 50°52’07.9” W, altitude 926 meters a.s.l, L.E.K. Lanés & R. S. Godoy & M. Fogaça, 30 Aug 2013; MCP 54455, 2 males, 28.0– 34.7 mm SL (1 C&S), 1 female, 25.6 (C&S), 28°22’47.9” S, 50°52’07.9” W, altitude 926 meters a.s.l., L.E.K. Lanés & A.B. Moraes, 10 May 2013; MCP 54456, 1 male, 34.8 mm SL (C&S), 1 female, 31.8 (C&S) 28° 22’43.6” S, 50°52’21.8” W, altitude 934 meters a.s.l., L.E.K. Lanés & A.B. Moraes, 10 May 2013. Diagnosis. Austrolebias botocudo differs from its congeners allocated in the subgenus Acrolebias by a unique male colour pattern consisting of body flank lilac grey with bluish white vertical bands (vs. flank pale yellowish brown, with dark brown bars in A. araucarianus; flank golden with purplish grey bars in A. carvalhoi; and flank dark bluish brown with yellowish golden or light grey vertical bands in A. nubium). The new species additionally differs from A. araucarianus and A. carvalhoi by males presenting melanophores irregularly distributed across the body (vs. melanophores absent in A. araucarianus and A. carvalhoi). Austrolebias botocudo differs from the remaining species of the subgenus Acrolebias, except from A. nubium, by presenting contact organs on anal fin in males (vs. contact organs absent). Austrolebias botocudo differ from A. nubium by the presence of discrete contact organs in the flank (vs. conspicuous contact organs), presence of generally only one contact organ by scale of lateral line (vs. two prominent contact organs by scale of lateral line), by presenting jaws slightly prognathous (vs. jaws short); by dorsal profile of head slight concave (vs. nearly convex), more neuromasts in the preopercular + mandibular series (36–43 vs. 26–35), a lower body depth in females (28.2–32.3 % SL vs. 32.6–37.8 % SL), and by presenting basihyal cartilage about 40–50% of basihyal length (vs. 60–70% of basihyal length). The new species differ from A. araucarianus by presenting neuromasts of the supraorbital series united (vs. first three neuromasts of the supraorbital series separated from the remaining neuromasts), preopercular and mandibular series of neuromasts united (vs. separated), by presenting pelvic fin well-developed (vs. pelvic-fin girdle rudimentary or absent), fewer scales in the longitudinal series (27–30 vs. 31–33), and by presenting more neuromasts in the ventral opercular series (2–4 vs. 1). Austrolebias botocudo differ from A. carvalhoi by presenting more neuromasts in the supraorbital (2–3+22–27 vs. 3+20) and lateral mandibular series (5–9 vs. 4), preopercular and mandibular series of neuromasts united (vs. separated), and by presenting basihyal cartilage about 40–50% of basihyal length (vs. 65% of basihyal length). Description. Morphometric data are presented in Table 1. Largest male examined 55.6 mm SL; largest female examined 39.3 mm SL. Dorsal profile slightly concave on head, convex from nape to the end of dorsal-fin base, nearly straight on caudal peduncle. Ventral profile convex from lower jaw to end of anal-fin base, about straight on caudal peduncle. Body moderately deep and compressed. Greatest body depth at vertical of pelvic fin origin. Lower jaw relatively long and prognathous, snout blunt. Dorsal-fin rays in males 22–24; in females 18–20. Dorsal fin sub-rectangular in males, with posterior tip rounded. Dorsal fin semi-circular in females. Origin of dorsal fin in males at vertical through neural spine of 7–9th vertebrae; in females through 10–11th vertebrae. Anal-fin rays in males 21–23, in females 18–20. Anal-fin sub-rectangular, with tip rounded in males. Anal-fin sub-triangular in females. Anteromedian rays of anal fin lengthened in females. Anal-fin origin at vertical through base of 3–5th dorsal-fin ray in males and through base of 2–4th dorsal-fin ray in females, and at vertical through pleural ribs of 8–9th vertebrae in males; and through 10–12th vertebrae in females. Caudal fin distal margin rounded, 24–27 rays in both sexes. Caudal-fin rays supported by last 4 vertebrae. Pectoral fin rays 12–14 in both sexes; pectoral-fin margin rounded. Pectoral-fin posterior tip reaching from before pelvic fin origin to 2nd anal-fin ray in males, from origin of pelvic fin or before it in females. Pelvic fins usually well-developed, with 5–6 rays, its tip extending from urogenital papilla to 2nd anal-fin ray in males, and to urogenital papilla in females. Urogenital papilla not attached to anal fin. Pelvic-fin bases medially separated by short interspace. Scales large and cycloid. Head and trunk entirely scaled, except ventral surface of head. Frontal squamation F, G, or H, sometimes irregularly arranged. No scales on dorsal and anal fin bases. Longitudinal series of scales 27–30, scales regularly arranged; transverse series of scales 11–16; 18–22 scale rows around caudal peduncle. Few, scattered contact organs on scales throughout body in males, when present located mainly at mid-ventral region of the flank from immediately distal tip of opercle to caudal peduncle. Usually one contact organ per scale of longitudinal series; rarely two contact organs along some scales of longitudinal series (present in only three males). Contact organs on distal portion of anteriormost 1–4 anal fin rays, present in 25% of males. Contact organs on outer region of 1–4 uppermost pectoral fin rays in males. Cephalic neuromasts: supraorbital 14–23, parietal 1–3, anterior rostral 1–2, posterior rostral 1, infraorbital 2–3+22–27, preorbital 2–3, otic 2–4, post-otic 2–5, supratemporal 1–2, median opercular 1, ventral opercular 2–4, preopercular + mandibular 36–43, lateral mandibular 5–9, paramandibular 1. Two neuromasts on caudal fin base. Six branchiostegal rays. Dermosphenotic ossification absent. Urohyal deep. Total number of vertebrae 28–29. Gill rakers in first branchial arch 3+8–9. Basihyal subtriangular, width about 60–75% of length; basihyal cartilage about 40–50% of total basihyal length. One to three teeth on second pharyngobranchial. Vomerine teeth absent. Colouration in life. Males (Fig. 2A–D). Ground colour of body lilac grey, brown in dorsal region, with 7–12 bluish white vertical bands, usually with narrower, overlapping or poorly defined stripes between well-defined bars. Some specimens in a dominant situation shown a darker coloured pattern, composed of a blue-purple background with vertical grey-green stripes. Pectoral and ventral region greyish. Melanophores irregularly distributed over the body, forming black dots, spots or irregular marks. Some specimens exhibit a concentration of melanophores in the middle of the flank, forming irregular black vertical bands. This pattern is more frequent in senile individuals. Opercle and preopercle greenish blue. Supraorbital and suborbital bars black. Pectoral fin hyaline, margin black. Pelvic fins dark blue or dark grey. Iris orange; dark vertical band crossing the eye. Dorsal fin dark blue with light grey elongated dots. Anal fin bluish-green with light, elongated grey dots; distal margin of anal fin darker. Caudal fin bluish grey with light grey dots and spots, distal margin hyaline. Females (Fig. 2E–H). Ground colour of body pale yellowish brown, generally with brown, dark brown or black spots distributed over the flank. Some individuals presenting a spotted pattern with several circular marks, which extends from dorsum to the medium flank. Dorsum generally pale yellowish brown. Some specimens present black melanophores in the dorsal region. Venter light yellowish grey. Sides of head yellowish brown, opercular region pale golden or bluish golden. Suborbital bar grey. Iris light yellow with transverse grey bar through centre of eye. Unpaired fins yellowish hyaline, with sparse and little conspicuous grey dots. Pectoral fin hyaline. Pelvic fin hyaline. Distribution. The species was recorded only in two temporary pools located in a highland grassland landscape at the rio Socorro, a tributary of upper rio Apuaê-Inhandava (upper rio Uruguay basin), in altitudes between 926 and 934 meters a.s.l., Vacaria municipality, Rio Grande do Sul State, southern Brazil (Fig. 3). Etymology. The specific epithet “botocudo” is a reference to an indigenous people, which formerly inhabited the region and belonged to the Xokleng Laklãnõ ethnic group, known during the 19th century as “Botocudos”, because adult men used a botoque (a plug-like wooden disk) to enlarge the lower lip. An allusion to a feature presented by the species, where the lower jaw seems more prominent than other species belonging to the subgenus Acrolebias. Ecological notes. Austrolebias botocudo was found only in two isolated high-altitude temporary wetlands, located within a private farmland, with size areas of 2,500 and 13,000 m 2. These two biotopes present clear (sometimes turbid), still water, with a latosol soil type, and are located in a grassland landscape (Fig. 4). The wetlands are close to each other (less than one kilometer apart) but are hydrologically isolated. A portion of the larger temporary pool was excavated, probably to become a water source for cattle. There are small intermittent drainages associated with the pools. The vegetation is composed mainly by stands of Peruvian watergrass (Luziola peruviana), peatbog (Sphagnum sp.), and other emergent, floating and submerse macrophytes. During winter and early spring, when the temporary pools present water, the minimum air temperature reaches 0°C (often negative temperatures are recorded), and frost, dew, fog and even snow can occur. The area where the biotopes inhabited by A. botucudo lie within are used for extensive cattle ranching. In the surroundings, mainly soybean crops and orchards (apples and pears) have drastically altered the natural high-altitude grasslands landscape. Conservation. In accordance with IUCN criteria, Austrolebias botocudo should be considered as “Critically Endangered”, with the criteria CR B2ab (ii, iii). The species presents an area of occupancy of 15,500 m 2 (AOO less than 10 km ²), present severely fragmented populations (a) with continued decline (b) in area of occupancy (ii) and quality of habitat (iii).

Sisyphus manni Montreuil, 2015 (Fig. 2E). Montreuil 2015c: 2���3 Type locality: Limpopo, Kruger National Park. Size: Male: length: 5.5���4.0 mm; width: 2.5���2.0 mm. Female: length: 5.5���4.0 mm, width 2.5���2.0 mm. Diagnosis: In S. manni, proximal elytral setae are primarily distributed uniformly becoming arranged in sparse tufts mid-basally and posteriorly, unlike in S. perissinottoi where dense tufts of setae occur across the entire elytra. S. manni bears three depressions on the pronotal disc, which are lacking in S. perissinottoi. Examined type material Holotype: (MHNH, through photograph without locality data). Examined non-type material: See Supplementary information. Distribution: S. manni is only known from South Africa. The species is associated with upland to highland grassland and open woodland in Gauteng (cited as S. alveatus by Davis et al. 2005), KwaZulu Natal, Mpumalanga and Northwest Provinces. It has also been collected in unshaded riverine vegetation in Limpopo (Fig. 4)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 15-16, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Montreuil, O. (2015 c) Premiers cas de brachypterisme dans le genre Sisyphus Latreille (Coleoptera, Scarabaeidae, Sisyphini). Annales de la Societe entomologique de France, 51 (4): 281 - 293.","Davis, A. L. V., Scholtz, C. H., & Deschodt, C. (2005) A dung beetle survey of selected Gauteng nature reserves: implications for conservation of the provincial scarabaeine fauna. African entomology, 13 (1): 1 - 16."]}

Sisyphus costatus (Thunberg, 1818) (Figs. 11, 12B). Peringu��y 1901:100; Gillet 1912:22; Arrow 1927:458; Haaf 1955:355���6; Ferreira 1972: 794���5; Harold 1869:984 Ateuchus costatus Thunberg, 1818 Thunberg 1818:412 Size: Male: length: 5.3��� 4.8 mm, width: 3.0���2.0 mm. Female: length: 5.3��� 4.2 mm, width: 2.6��� 2 mm. Type locality: (Cape of Good Hope) South Africa Diagnosis: S. costatus is close in appearance to S. australis sp. n. In both species, the elytral interstriae bear alternating rows of well-developed or less-developed setae (Fig. 11F). However, S. costatus is distinguished by an upcurved anterior clypeal edge and a concavity between the medial clypeal teeth. In addition, it lacks a carina between the clypeus and the frons (Fig. 11G). Examined type material (1♀ SMNH) Lectotype (designated here): Red label / Uppsala University Zoology Museum; Thunberg, saml.nr. 3206., Ateuchus costatus, Cap. minutus. F. TYP/ / Lectotype, det. GM Daniel, 2016 /. Examined non-type material: See Supplementary information. Distribution: S. costatus has been recorded in moist upland savanna and highland grassland (Davis et al. 1999, Davis et al. 2005) from the Eastern Cape to the north of South Africa (Fig. 13). Remarks: The species was described by Thunberg (1818) as Ateuchus costatus. Gillet (1912) transferred Thunberg���s species to the genus Sisyphus., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 29-30, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Thunberg, C. P. (1818) Coleoptera Capensia, Antennis Lamellatis, sive, clava fissili instructa, descripta, memoires de l'Acad. T. VI.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Davis, A. L. V., Scholtz, C. H., & Chown, S. L. (1999) Species turnover, community boundaries and biogeographical composition of dung beetle assemblages across an altitudinal gradient in South Africa. Journal of Biogeography, 26 (5): 1039 - 1055.","Davis, A. L. V., Scholtz, C. H., & Deschodt, C. (2005) A dung beetle survey of selected Gauteng nature reserves: implications for conservation of the provincial scarabaeine fauna. African entomology, 13 (1): 1 - 16."]}

Sisyphus caffer Boheman, 1857 (Figs. 5, 6B). Boheman 1857: 195���6; Harold 1869: 984; P��ringuey 1902: 106���7; Haaf 1955: 362���4; Ferreira 1972: 82, Sch��fer & Fischer 1992:121; Sch��fer & Fischer 2001:53 Size: Male: length: 7.8��� 6.9 mm, width: 4.4��� 3.5 mm. Female: length: 7.6��� 7.1 mm, width: 4.5���3.0 mm. Type locality: Caffraria (South Africa). Diagnosis: S. caffer is close in appearance to S. sordidus Boheman, 1857. However, they can be separated morphologically by the following features: S. caffer bears much finer elytral setae, which are distributed in single rows; S. sordidus bears thicker elytral setae, which are distributed in non-single rows forming bunches. In S. caffer, pronotal setae are inserted between ocellate punctures (Fig. 5F), whereas in S. sordidus, the setae originate from the centre of ocellate punctures (Fig. 5H). Furthermore, S. sordidus occurs in unshaded lowland vegetation associated with warm temperatures whereas S. caffer is associated with highland grassland under cooler conditions. Examined type material Holotype (photograph): (1♂ NHRS): /type/; /Caffraria/; /leg: J.Wahlb /; /309; 72/; /244; 73/; / S. caffer Bohm /; red label: /typus/; white label (handwritten): /TYPE Sisyphus caffer BOHEMAN, det. J. Ferrer /; /Naturhistoriska, Riksmuseet Stockholm; loan 37/90/; /9385 E92+/; /NHRS-SRAH 000000221/. Examined non-type material: See Supplementary information. Distribution: S. caffer occurs in South Africa, Namibia, Lesotho and Bostwana. The species has been recorded in upland and highland grasslands under cooler conditions (Davis et. al. 2005) in both South Africa and Lesotho (29.7��S 27.4��E). In South Africa, the localities extend northwards from the Eastern Cape and eastern Free State to the latitude of Pretoria (28.19��E 25.80��S) and also along the eastern escarpment of KwaZulu Natal and Mpumalanga. The lowlands beyond the northern Gauteng Mountains of the Witwatersrand (24.54��S 27.08��E) may act as barriers, although the species is also found in isolated uplands further to the northwest, such as the Blouberg. S. caffer has also been recorded in unshaded vegetation at the margins of its range in Kneneng (Botswana) (23.78��S 25.27��E) and in a dry upland outlier in the north of Namibia (Kaoko Otavi) (Fig. 7). Kaoko Otavi represents a disjunction in the distribution of S. caffer from its main range on the Highveld of South Africa. It is noteworthy that N. macrorubrus and Proagoderus lanista (Castelnau, 1840) also show similar disjunct occurrences on the South African Highveld and as rarities in north Namibia (Davis 1997). Remarks: We cannot confirm or refute the validity of S. crispatus because the type specimen has been lost. Furthermore, the original description and image presented by Gory (1833) are imprecise and insufficient to determine the identity of the species. S. crispatus was described from the same imprecise type locality (Caffraria) as two other species (S. caffer and S. sordidus). Under these circumstances, we are not confident to state which of the two species might be a junior synonym of S. crispatus. According to the ICZN, article 75.5 ���when an author considers that the taxonomic identity of a nominal species-group taxon cannot be determined from its existing name-bearing type, its name is a nomen dubium ���. Therefore, we propose S. crispatus as a nomen dubium., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 20-21, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Boheman, C. H. (1857) Insecta Caffrariae annis 1838 - 1845 a JA Wahlberg collecta. Pars. II. Coleoptera (Scarabaeides). Holmiae: Offieina Nordstedtiana. - 395 pp, 1.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Schafer, P., & Fischer, E. (1992) Zur Scarabaeiden-Fauna Rwandas, Zentralafrika. Sisyphini (Insecta: Coleoptera: Lamellicornia). Senckenbergiana biologica, 72 (1 - 3), 119 - 137.","Schafer, P. & Fischer, E. (2001) Zur Scarabaeidenfauna Rwandas, Zentralafrika 2, Gymnopleurini und nachtrag Sisyphini, Mitteilungen des internationalen entomologischen Vereins supplement 9: 1 - 93.","Davis, A. L. V. (1997) Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.). african Journal of ecology, 35 (1): 10 - 38.","Gory, M. (1833) Monographie du genre Sisyphe. Mequignon-Marvis pere et fils, Paris. 1 - 15 p."]}

Neosisyphus kuehni (Haaf, 1955) (Figs. 26, 27B) Montreuil 2015: 28. Sisyphus kuehni Haaf, 1955 Haaf 1955: 345; Montreuil 2015: 28. Type locality: Natal (Griqualand), South Africa. Size: Male: length: 13.5���10; width: 6.5���5.5; Female: length: 13.0���11; width: 6.5���5.6. Diagnosis: N. kuehni bears a gibbosity apically on the metafemur, which is lacking in the closely-related N. quadricollis. The projecting metatrochanter in males of N. kuehni is also longer, comprising over half of the total length of the metafemur. Unlike in N. quadricollis, the elytra in N. kuehni are not laterally emarginate. Examined non-type material: See Supplementary information. Distribution: N. kuehni occurs in highland grassland under the moist to wet conditions of the eastern escarpment of South Africa (Eastern Cape, KwaZulu Natal and Mpumalanga) (Fig. 28). Conservation status: N. kuehni was assessed as a Data Deficient species for the IUCN Red Date List (Davis 2013g). Further collecting and monitoring is required to accurately assess the conservation status of this species., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 53-54, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Davis, A. L. V. (2013 g) Neosisyphus kuhni. The IUCN Red List of Threatened Species 2013: e. T 138156 A 537143. http: // dx. doi. org / 10.2305 / IUCN. UK. 20132. RLTS. T 138156 A 537143. en. (accessed 14 October 2016)."]}

Gyronotus dracomontanus Deschodt & Davis, new species (Figs. 1C, 3) Type material. Holotype, female: ���Z.A.29 | Bergville Dist. | Drakensberg || Prairie subalpine | Alt. 2,400m., x-1960 || N. Leleup Leg. ��� (TMSA). Paratypes: two females, no locality data: ���ex collection of | Pietermaritzburg | Museum ��� (UPSA). Description of the holotype, female. Size. Length 12.0 mm and width 8.0 mm with head extended (not deflexed). Habitus. Body shape ovoid. Black in appearance. Head. Tip of clypeus bidentate, separated by a wide ���U��� shaped depression. Surface with dense punctation bearing very short fine setae almost touching each other. Clypeogenal sutures fairly distinct and nearly straight. Frontogenal suture clearly visible only at external one tenth; middle four fifths not visible. Small round tubercle medially on frontogenal suture. Antennae dark orange. Thorax. Pronotal disc densely punctate, punctures round to slightly ovoid, mostly touching one another. Lateral margin with obtuse angle almost halfway, otherwise relatively straight. Surfaces of all thoracic sternites shagreened between punctation. Prosternum with medium sized punctures that are separated by about half a puncture diameter. Mesosternum with medium sized punctures that are close together frontally and further apart anteriorly; row of overlapping setigerous punctures on mesometasternal suture. Metasternum with small sized punctures spaced up to three diameters apart. Mesometasternal suture clear, evenly slightly curved. Elytra. Interstriae punctate, punctures small and setigerous separated by half up to two puncture diameters. Surface otherwise shagreened appearing slightly rugose under low magnification (x10). Striae indistinct, visible as slight impressions bearing small punctures that are spaced one to two puncture diameters apart. Pygidium. Faintly punctate, punctures small and setigerous about one to three puncture diameters apart, laying closer together dorsally and further apart ventrally, surface otherwise shagreened giving a dull appearance. Distribution. Gyronotus dracomontanus Deschodt & Davis, new species most probably occurs in the high grassland above 2000 m between Royal Natal National Park [S28.702�� E28.915��] and Cathedral Peak [S28.925�� E29.134��]. Etymology. This species is named for the Drakensberg mountain range where it occurs. Diagnosis. This species can be separated from most other Gyronotus species by having prominent, dense setae on the elytra. The highland grassland species, Gyronotus dracomontanus Deschodt & Davis, new species, has six or less adjacent setae on the second interval of the interstriae compared to eight or more in the coastal forest species, Gyronotus pumilus., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2019, New southern African species and a revalidation in the dung beetle genus Gyronotus van Lansberge, 1874 (Scarabaeidae: Scarabaeinae) with an updated key, pp. 275-282 in Zootaxa 4624 (2) on pages 276-277, DOI: 10.11646/zootaxa.4624.2.10, http://zenodo.org/record/3259222

443. San Javier's Pygmy Rice Rat Oligoryzomys brendae French: Colilargo de Brenda / German: San-JavierZwergreisratte / Spanish: Rata arrocera pigmea de San Javier Other common names: Brenda's Colilargo, San Javier Colilargo Taxonomy. Oligoryzomys brendae Massoia, 1998, Cerro San Javier, 1000 m, Tucuman, Argentina. Most available literature refers to O. brendae as O. longicaudatus. Monotypic. Distribution. Highlands from Jujuy and Salta to La Rioja provinces, NW Argentina; it may occur in S Bolivia, limits unresolved. Descriptive notes. Head—body 78-101 mm (mean 90 mm), tail 112-135 mm, ear 17-20 mm, hindfoot 24-28 mm; weight 12-34 g. San Javier's Pygmy Rice Rat is large, with orangish brown dorsum and somewhat grizzled appearance; ventral hairs are basally gray and whitish to ocherous at tips; ears are short, rounded, and dark brown;tail is longer than head-body length and sharply bicolored; skull is relatively robust, with short and broad rostrum, well-expanded zygomatic arches, and inflated and broad braincase; interorbital constriction is narrow and hourglass-shaped, with slightly defined supraorbital ridges; and incisive foramina are relatively long, posteriorly extended to anterior border of M'. Chromosomal cmplement is 2n = 58, FN = 74. Habitat. Forested and highland grassland environments of Yungas at elevations of 700-2900 m. San Javier's Pygmy Rice Rat was restricted to isolated humid ravines in xeric Dry Chaco and Monte Desert formations. Food and Feeding. One San Javier's Pygmy Rice Rat had insects and vegetation in its stomach. Breeding. Non-reproductive males and females were reported in dry season (May-September). Young were found in dry season, suggesting that most reproduction occurred in wet season (November—February). Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Other than deforestation, no major threats apparently face San Javier’s Pygmy Rice Rat, but there is no information on its abundance and population trends. Bibliography. Crespo (1974), Diaz et al. (1997, 2000), Ferro & Barquez (2017), Gamboa et al. (2016), Gonzalez-Ittig et al. (2014), Jayat & Ortiz (2010), Jayat & Pardinas (2008a), Mares, Ojeda & Barquez (1989), Mares, Ojeda, Braun et al. (1997), Ojeda & Mares (1989), Ojeda, Barquez et al. (2008), Sandoval et al. (2015), Teta, Jayat et al. (2013), Weksler & Bonvicino (2015b)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Cricetidae, pp. 204-535 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 436, DOI: 10.5281/zenodo.6707142

Nata erugata sp. nov. Figures 4 C, 6 C, 29–32 Natalina caffrula — Haas, 1936: 18 (in part, Mkuzi). Nata vernicosa — Bruggen, 1966: 376, fig. 54. Bruggen, 1969: 37. Bruggen, 1970 a: 468 (in part). Bruggen, 1970 b: 59. Bruggen, 1985: 286 (in part). Etymology. From erugo (L.)—to clear of wrinkles, erugatus (L.)–without wrinkles, smooth. Type material. Holotype (Fig. 30 A–C): SOUTH AFRICA: KWAZULU-NATAL: Monk’s Cowl area (29.05479 °S: 29.38287 °E), 1625 m, indigenous forest, in leaf-litter, F 78, M. Hamer et al., 16 /iii/ 2006 (NMSA W 5599 /T 3906). Paratypes (all NMSA unless indicated otherwise): SOUTH AFRICA: KWAZULU-NATAL: Pongola Bush Nat. Res. (27.323 °S: 30.4817 °E), 1300 m, montane Podocarpus forest, in leaf-litter and under logs, D. Herbert & NPB, 19 /i/ 1995 (V 776 /T 3915, one adult and three juvenile specimens; ELM D 18003 /T031, one specimen); Majuba area (27.477 °S: 29.685 °E), 1700 m, montane Podocarpus forest, in leaf-litter, D. Herbert, 28 /i/ 1998 (V 5932 /T 3909, one adult and three juvenile specimens); De Beers farm, approx. 15 km NE of Van Reenen (28.30403 °S: 29.50240 °E), 1841 m, afrotemperate forest, in leaf-litter, A. Moussalli, D. Stuart-Fox & M. Cunningham, 9 /x/ 2006 (W 4688 / T 3910, one specimen); Van Reenen (28.405 °S; 29.390 °E), iii/ 1918 (A 7069 /T 3911, five specimens; ELM D 18002 / T030, one specimen); Kranskop (28.967 °S: 30.863 °E), Falcon coll’n, i/ 1922 (NMSA A 7070 /T 3912, three specimens). FREE STATE: Platberg Nat. Res. (28.27631 °S: 29.20772 °E), 2100 m, montane shrubland/grassland, in leaf-litter, A. Moussalli, D. Stuart-Fox & M. Cunningham, 11 /xii/ 2006 (W 5621 /T 3908, two specimens); Golden Gate Highlands National Park (28.49430 °S: 28.61868 °E), 2000 m, gully thicket/grassland, buried in shrubs, A. Moussalli & D. Stuart-Fox, 28 /xii/ 2005 (W 4243 /T 3913, one specimen); Zastron area, Mapaya Hill (30.27439 °S: 27.21902 °E), ~ 1860 m, in soil amongst boulders in narrow, shaded cleft above sandstone cliff, st'n 08-059, D. Herbert & L. Davis, 22 /xi/ 2008 (W 6573 /T 3907, three specimens). E. CAPE: Witteberge, Balloch farm, north of Barkly East (30.71255 °S: 27.69507 °E), Lesotho highland grassland, under rock, D. Stuart-Fox, 1 /iv/ 2005 (W 4836 / T 3914, one adult and two juvenile specimens). Additional material examined: See Appendix 4. Identification. Differs from Nata vernicosa in that the apical surface is almost completely devoid of regular, close-set, axial riblets; although axial sculpture remains, it is present only in the form of weak, irregularly spaced growth-lines. More similar to Nata watsoni sp. nov., but that species has very different radula morphology and occurs in a different region and biome (Albany Thicket). Shell (Fig. 30): Lenticular, occasionally discoidal; whorls comparatively shallow and spire often low; largest specimens comprising 3.25–3.5 teleoconch whorls; periphery at or more usually slightly below mid-whorl; suture above periphery, at most weakly descendant prior to aperture. Protoconch diameter 1.0– 1.9 mm, comprising ±1.0 whorl, smooth and glossy, junction with teleoconch usually rather weakly defined. Apical surface of teleoconch smooth and glossy, lacking regular axial riblets and sculptured only by weak growth irregularities; these strongest below suture, weakest at periphery and on base; no significant spiral sculpture, but some specimens with faint traces of vermiform wrinkles in places. Base and umbilical margin evenly rounded; umbilicus of moderate width, steep-sided; growth-lines slightly stronger within umbilicus. Aperture roundly or distinctly ovate, often obliquely descendant (Fig. 30 J); peristome interrupted by bulging parietal region; outer lip thin with membranous periostracal fringe (commonly lost); outer lip shallowly sinuous when undamaged; upper part of columella lip extending forward and slightly reflected. Shell translucent milky-white when fresh; periostracum glossy, pale yellowish-brown to honey-gold, commonly with some axial variation in intensity, producing unevenly spaced, slightly darker axial banding, but this generally weak. Dimensions: Holotype, diameter 15.5 mm, height 8.1 mm; largest specimen, diameter 21.0 mm, height 11.4 mm; H:D of adults 0.46–0.60 [N= 34]. Living animal (Figs 4 C, 31): Body generally pale; head and neck greyish-white to cream, dorsal region of neck sometimes pale grey or pale bluish-grey; optic tentacles a slightly darker shade of grey; foot paler beneath shell; posterior portion of foot relatively short; some individuals with microscopic orange pigment granules in skin, particularly on hind end of foot; skin texture moderately well defined; mantle edge pale, also with minute orange granules in some individuals; lining of mantle cavity with little or no dark pigmentation, except for a diffuse blackish mark just inside pneumostome, visible through shell; kidney trigonal, pale buff, not conspicuous through shell. Radula (Figs 6 C, 32): Very similar to that of Nata vernicosa, but usually with fewer lateral teeth. Formula 0+(7–8)+(3–4); length up to 12.0 mm, with 24–33 V-shaped rows of teeth; teeth progressively larger from anterior to posterior of radula; rachidian absent; largest tooth (number 7 or 8) taken to be outermost lateral. Tooth morphology as in N. vernicosa. Distal genitalia: As in Nata vernicosa. Epiphallus enters penis between one-third and two-thirds length of the latter from its base; internally, lining of penis forms a raised rim around epiphallus pore; basal portion of bursa copulatrix duct swollen (particularly so in specimens from Hogsback), tapering to mid region and evenly slender thereafter, but constricted prior to bursa copulatrix which is small, subovate to reniform and thin walled. Distribution (Fig. 29): Endemic to eastern South Africa and southern Mozambique, but relatively widely distributed within this region, ranging from Hogsback in the south west, through western Lesotho, eastern Free State and Gauteng, into western and northern KwaZulu-Natal, the Lowveld of Mpumalanga and southern Mozambique. Throughout the southern and western parts of its range the species is largely restricted to altitudes over 1000 m and extending to 2200 m along the KwaZulu-Natal escarpment edge. However, the populations in north-eastern KwaZulu-Natal (Zululand), the Mpumalanga Lowveld (southern Kruger National Park) and Mozambique occur at much lower altitudes (30–300 m). Habitat. At altitudes above 1000 m the species occurs primarily in afrotemperate forest and more thickly vegetated habitats such as Leucosidea scrub. However, specimens have also been found in sheltering microhabitats in more open vegetation, even montane grassland. At lower altitudes in Zululand and the Kruger National Park it occurs in a variety of vegetation types within the Savannah Biome. Remarks. When discussing Pfeiffer’s ‘ var. minor ’, Connolly (1939) noted that although Pfeiffer’s description ‘ costulis superficiei obsoletioribus ’ (Pfeiffer 1853: 95) may apply to an immature example of Nata liparoxantha, ‘…there extends from the Cape Province to Delagoa a hitherto undetermined race, which may represent his variety. It differs from vernicosa solely in being very much smoother, the sculpture consisting of fine growth striolae instead of costulae’ (Connolly 1939: 100). We have likewise been able to discern such a ‘race’, but we do not consider that it can be considered conspecific with Pfeiffer’s var. minor. Examination of the lectotype of the latter at the SMNH (see above regarding its authenticity) reveals that although the axial sculpture is weak, distinct and regularly spaced axial riblets are nonetheless present on the adapical surface and the specimen is simply a weakly sculptured example of the variable N. vernicosa. Since all the other names currently considered to be junior synonyms of N. vernicosa likewise relate to material with distinct axial riblets, we are therefore obliged to propose a new name for the present material. Throughout much of its distribution Nata erugata sp. nov. is the only species of Nata to be found, but at the lower limits of its altitudinal range, particularly in E. Cape and KwaZulu-Natal, there is some geographical overlap with Nata vernicosa (Fig. 29). However, further studies are needed to establish whether and to what extent the two species co-occur. There is certainly evidence that this may be the case at some localities (Hogsback, Umthatha area, Karkloof and Pilgrim’s Rest area). The occurrence of populations of what is largely a higher altitude species in the hotter more low-lying regions of northern KwaZulu-Natal, the Kruger National Park and southern Mozambique (Fig. 29), at altitudes of well below 1000 m, is anomalous and hard to explain. It is certainly suggestive of there being a further cryptic lineage in what we group under one species. For this reason, we have not included any specimens from these localities in the type material for this species. This is an issue needing to be studied in more detail using molecular data. Conservation. As interpreted here, Nata erugata sp. nov. is a widespread and moderately common species occurring in a range of habitat types. As such, its conservation is not at present a matter of concern.

Nata erugata sp. nov. Figures 4 C, 6 C, 29–32 Natalina caffrula — Haas, 1936: 18 (in part, Mkuzi). Nata vernicosa — Bruggen, 1966: 376, fig. 54. Bruggen, 1969: 37. Bruggen, 1970 a: 468 (in part). Bruggen, 1970 b: 59. Bruggen, 1985: 286 (in part). Etymology. From erugo (L.)—to clear of wrinkles, erugatus (L.)–without wrinkles, smooth. Type material. Holotype (Fig. 30 A–C): SOUTH AFRICA: KWAZULU-NATAL: Monk’s Cowl area (29.05479 °S: 29.38287 °E), 1625 m, indigenous forest, in leaf-litter, F 78, M. Hamer et al., 16 /iii/ 2006 (NMSA W 5599 /T 3906). Paratypes (all NMSA unless indicated otherwise): SOUTH AFRICA: KWAZULU-NATAL: Pongola Bush Nat. Res. (27.323 °S: 30.4817 °E), 1300 m, montane Podocarpus forest, in leaf-litter and under logs, D. Herbert & NPB, 19 /i/ 1995 (V 776 /T 3915, one adult and three juvenile specimens; ELM D 18003 /T031, one specimen); Majuba area (27.477 °S: 29.685 °E), 1700 m, montane Podocarpus forest, in leaf-litter, D. Herbert, 28 /i/ 1998 (V 5932 /T 3909, one adult and three juvenile specimens); De Beers farm, approx. 15 km NE of Van Reenen (28.30403 °S: 29.50240 °E), 1841 m, afrotemperate forest, in leaf-litter, A. Moussalli, D. Stuart-Fox & M. Cunningham, 9 /x/ 2006 (W 4688 / T 3910, one specimen); Van Reenen (28.405 °S; 29.390 °E), iii/ 1918 (A 7069 /T 3911, five specimens; ELM D 18002 / T030, one specimen); Kranskop (28.967 °S: 30.863 °E), Falcon coll’n, i/ 1922 (NMSA A 7070 /T 3912, three specimens). FREE STATE: Platberg Nat. Res. (28.27631 °S: 29.20772 °E), 2100 m, montane shrubland/grassland, in leaf-litter, A. Moussalli, D. Stuart-Fox & M. Cunningham, 11 /xii/ 2006 (W 5621 /T 3908, two specimens); Golden Gate Highlands National Park (28.49430 °S: 28.61868 °E), 2000 m, gully thicket/grassland, buried in shrubs, A. Moussalli & D. Stuart-Fox, 28 /xii/ 2005 (W 4243 /T 3913, one specimen); Zastron area, Mapaya Hill (30.27439 °S: 27.21902 °E), ~ 1860 m, in soil amongst boulders in narrow, shaded cleft above sandstone cliff, st'n 08-059, D. Herbert & L. Davis, 22 /xi/ 2008 (W 6573 /T 3907, three specimens). E. CAPE: Witteberge, Balloch farm, north of Barkly East (30.71255 °S: 27.69507 °E), Lesotho highland grassland, under rock, D. Stuart-Fox, 1 /iv/ 2005 (W 4836 / T 3914, one adult and two juvenile specimens). Additional material examined: See Appendix 4. Identification. Differs from Nata vernicosa in that the apical surface is almost completely devoid of regular, close-set, axial riblets; although axial sculpture remains, it is present only in the form of weak, irregularly spaced growth-lines. More similar to Nata watsoni sp. nov., but that species has very different radula morphology and occurs in a different region and biome (Albany Thicket). Shell (Fig. 30): Lenticular, occasionally discoidal; whorls comparatively shallow and spire often low; largest specimens comprising 3.25–3.5 teleoconch whorls; periphery at or more usually slightly below mid-whorl; suture above periphery, at most weakly descendant prior to aperture. Protoconch diameter 1.0– 1.9 mm, comprising ±1.0 whorl, smooth and glossy, junction with teleoconch usually rather weakly defined. Apical surface of teleoconch smooth and glossy, lacking regular axial riblets and sculptured only by weak growth irregularities; these strongest below suture, weakest at periphery and on base; no significant spiral sculpture, but some specimens with faint traces of vermiform wrinkles in places. Base and umbilical margin evenly rounded; umbilicus of moderate width, steep-sided; growth-lines slightly stronger within umbilicus. Aperture roundly or distinctly ovate, often obliquely descendant (Fig. 30 J); peristome interrupted by bulging parietal region; outer lip thin with membranous periostracal fringe (commonly lost); outer lip shallowly sinuous when undamaged; upper part of columella lip extending forward and slightly reflected. Shell translucent milky-white when fresh; periostracum glossy, pale yellowish-brown to honey-gold, commonly with some axial variation in intensity, producing unevenly spaced, slightly darker axial banding, but this generally weak. Dimensions: Holotype, diameter 15.5 mm, height 8.1 mm; largest specimen, diameter 21.0 mm, height 11.4 mm; H:D of adults 0.46–0.60 [N= 34]. Living animal (Figs 4 C, 31): Body generally pale; head and neck greyish-white to cream, dorsal region of neck sometimes pale grey or pale bluish-grey; optic tentacles a slightly darker shade of grey; foot paler beneath shell; posterior portion of foot relatively short; some individuals with microscopic orange pigment granules in skin, particularly on hind end of foot; skin texture moderately well defined; mantle edge pale, also with minute orange granules in some individuals; lining of mantle cavity with little or no dark pigmentation, except for a diffuse blackish mark just inside pneumostome, visible through shell; kidney trigonal, pale buff, not conspicuous through shell. Radula (Figs 6 C, 32): Very similar to that of Nata vernicosa, but usually with fewer lateral teeth. Formula 0+(7–8)+(3–4); length up to 12.0 mm, with 24–33 V-shaped rows of teeth; teeth progressively larger from anterior to posterior of radula; rachidian absent; largest tooth (number 7 or 8) taken to be outermost lateral. Tooth morphology as in N. vernicosa. Distal genitalia: As in Nata vernicosa. Epiphallus enters penis between one-third and two-thirds length of the latter from its base; internally, lining of penis forms a raised rim around epiphallus pore; basal portion of bursa copulatrix duct swollen (particularly so in specimens from Hogsback), tapering to mid region and evenly slender thereafter, but constricted prior to bursa copulatrix which is small, subovate to reniform and thin walled. Distribution (Fig. 29): Endemic to eastern South Africa and southern Mozambique, but relatively widely distributed within this region, ranging from Hogsback in the south west, through western Lesotho, eastern Free State and Gauteng, into western and northern KwaZulu-Natal, the Lowveld of Mpumalanga and southern Mozambique. Throughout the southern and western parts of its range the species is largely restricted to altitudes over 1000 m and extending to 2200 m along the KwaZulu-Natal escarpment edge. However, the populations in north-eastern KwaZulu-Natal (Zululand), the Mpumalanga Lowveld (southern Kruger National Park) and Mozambique occur at much lower altitudes (30–300 m). Habitat. At altitudes above 1000 m the species occurs primarily in afrotemperate forest and more thickly vegetated habitats such as Leucosidea scrub. However, specimens have also been found in sheltering microhabitats in more open vegetation, even montane grassland. At lower altitudes in Zululand and the Kruger National Park it occurs in a variety of vegetation types within the Savannah Biome. Remarks. When discussing Pfeiffer’s ‘ var. minor ’, Connolly (1939) noted that although Pfeiffer’s description ‘ costulis superficiei obsoletioribus ’ (Pfeiffer 1853: 95) may apply to an immature example of Nata liparoxantha, ‘…there extends from the Cape Province to Delagoa a hitherto undetermined race, which may represent his variety. It differs from vernicosa solely in being very much smoother, the sculpture consisting of fine growth striolae instead of costulae’ (Connolly 1939: 100). We have likewise been able to discern such a ‘race’, but we do not consider that it can be considered conspecific with Pfeiffer’s var. minor. Examination of the lectotype of the latter at the SMNH (see above regarding its authenticity) reveals that although the axial sculpture is weak, distinct and regularly spaced axial riblets are nonetheless present on the adapical surface and the specimen is simply a weakly sculptured example of the variable N. vernicosa. Since all the other names currently considered to be junior synonyms of N. vernicosa likewise relate to material with distinct axial riblets, we are therefore obliged to propose a new name for the present material. Throughout much of its distribution Nata erugata sp. nov. is the only species of Nata to be found, but at the lower limits of its altitudinal range, particularly in E. Cape and KwaZulu-Natal, there is some geographical overlap with Nata vernicosa (Fig. 29). However, further studies are needed to establish whether and to what extent the two species co-occur. There is certainly evidence that this may be the case at some localities (Hogsback, Umthatha area, Karkloof and Pilgrim’s Rest area). The occurrence of populations of what is largely a higher altitude species in the hotter more low-lying regions of northern KwaZulu-Natal, the Kruger National Park and southern Mozambique (Fig. 29), at altitudes of well below 1000 m, is anomalous and hard to explain. It is certainly suggestive of there being a further cryptic lineage in what we group under one species. For this reason, we have not included any specimens from these localities in the type material for this species. This is an issue needing to be studied in more detail using molecular data. Conservation. As interpreted here, Nata erugata sp. nov. is a widespread and moderately common species occurring in a range of habitat types. As such, its conservation is not at present a matter of concern., Published as part of David G. Herbert & Adnan Mousalli, 2016, Revision of the dwarf cannibal snails (Nata s. l.) of southern Africa — Nata s. s. and Natella (Mollusca: Gastropoda: Rhytididae), with description of three new species, pp. 1-67 in Zootaxa 4094 (1) on pages 41-44, DOI: 10.11646/zootaxa.4094.1.1, http://zenodo.org/record/266544, {"references":["Haas, F. (1936) Binnen-Mollusken aus Inner-Afrika, hauptsachlich gesammelt von Dr. F. Haas wahrend der Schomburgk- Expedition in den Jahren 1931 / 32. Abhandlungen Senckenbergischen Naturforschenden Gesellschaft, 431, 1 - 156.","Bruggen, A. C. van (1966) The terrestrial Mollusca of the Kruger National Park: a contribution to the malacology of the Eastern Transvaal. Annals of the Natal Museum, 18, 315 - 399.","Bruggen, A. C. van (1969) Studies on the land molluscs of Zululand, with notes on the distribution of land molluscs in southern Africa. Zoologische Verhandelingen, 103, 1 - 116.","Bruggen, A. C. van (1970 a) Non-marine Mollusca. Chapter 18. In: Hanstrom, B., Brinck, P. & Rudebeck, G. (Eds.), South African Animal Life. Results of the Lund University Expedition in 1950 - 1951. Vol XIV. Statens Naturvetenskapliga Forskningsrad, Stockholm, pp. 445 - 476.","Bruggen, A. C. van (1970 b) The terrestrial Mollusca of the Mkuzi and Ndumu Game Reserves, Zululand. Lammergeyer, 11, 58 - 59.","Bruggen, A. C. van (1985) The terrestrial molluscs of Lesotho (southern Africa), a first contribution, with detailed notes on Archachatina machachensis (Mollusca, Gastropoda). Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Series C, 88, 267 - 296.","Connolly, M. (1939) A monographic survey of South African non-marine Mollusca. Annals of the South African Museum, 33, 1 - 660."]}

Leptanilloides chihuahuaensis (Figs. 1, 2) Holotype male: [United States] TEXAS, Jeff Davis Co., Davis Mts. St. Pk., 5118 ', 30 �� 35 ' 27 "N 103 �� 56 ' 26 "W, 22��� 25 July 2014, T.L. Schiefer, Malaise trap on hillside on highland grassland, W.H. Cross Expedition. [Holotype deposited in Harvard Museum of Comparative Zoology (MCZC), Cambridge, MA]. Specimen code MEM 207971. Paratype males: Two point-mounted specimens with the same data as the holotype but with unique specimen code: MEM 207972 and MEM 207973 [both specimens in Mississippi Entomological Museum (MEM), Mississippi State, MS, USA]; and three specimens, one point-mounted and two in 95 % ethanol, with the following collection data: [United States] TEXAS, Jeff Davis Co., Davis Mts. St. Pk., 4915 ', 30 �� 36 '08"N 103 �� 54 ' 55 "W, 21���25 July 2014, T.L. Schiefer, Malaise trap in riparian area in desert scrub/grassland, W.H. Cross Expedition with unique specimen codes: MEM 207974 [point mounted specimen deposited in MEM], MEM 207975 [preserved in 95 % ethanol for DNA study, deposited in MEM] and MEM 207976 [preserved in 95 % ethanol for DNA study, deposited in MEM]. Holotype measurements and indices: HW 0.24, HL 0.24, EL 0.11, MDL 0.13, SL 0.08, PedL 0.06, FlaLI 0.04, FlaLII 0.04, FlaLXI 0.12, MH 0.29, ML 0.46, PrW 0.20, PW 0.09, PL 0.07, AIIIW 0.14, AIIIL 0.07, AIVW 0.20, AIVL 0.09, FFeW 0.05, FFeL 0.23, HFeL 0.24, HTiL 0.26, FWL 1.46, CI 100, MI 63, PI 128. Additional paratype measurements and indices (two males): HW 0.24, HL 0.22, EL 0.09���0.11, MDL 0.13, SL 0.09, PedL 0.06���0.07, FlaLI 0.04, FlaLII 0.04, FlaLXI 0.11���0.12, MH 0.30���0.31, ML 0.45���0.46, PrW 0.18, PW 0.07���0.08, PL 0.06���0.07, AIIIW 0.15, AIIIL 0.07���0.10, AIVW 0.18���0.19, AIVL 0 0.10, FFeW 0.05, FFeL 0.21, HFeL 0.23, HTiL 0.26, FWL 1.38, CI 109, MI 65���69, PI 114���117. Etymology. Named for the Chihuahuan Desert region where specimens were collected. Diagnosis (male). Of the six males that have been described in the Leptanilloides genus-group, L. chihuahuaensis is most similar to " Leptanilloidinae male 1 " from southern Mexico described by Borowiec & Longino (2011). Leptanilloides chihuahuaensis differs from " Leptanilloidinae male 1 " by having finer and sparser pilosity, in dorsal view the petiole being widened anteriorly and posteriorly, parameres being narrower and incurved, and the hindwing having three hamuli (instead of two). Males of L. chihuahuaensis differ from males of Amyrmex golbachi Kusnezov, 1953 by lacking submarginal cells and a stigma in the forewing, possessing a free M vein, and having a larger paramere (about as long as petiole); from L. mckennae Longino 2003 by lacking submarginal cells and a stigma in forewing and having a much shorter paramere (in side view) (Ward 2007); from L. nubecula Donaso, Vieira & Wild, 2006 by being light to medium brown instead of dark brownish-black and being much smaller (HL 0.22���0.24 in L. chihuahuaensis vs. HL 0.32 in L. nubecula), and from both " Leptanilloidinae male 2 and male 3 " (Borowiec & Longino 2011) by the much smaller size (HL 0.22���0.24 vs. HL 0.30���0.33 of " Leptanilloidinae male 2 and male 3 ") and by lacking submarginal cells and stigma in the forewing. Five other male morphotypes unassociated with workers were briefly discussed by Ward & Brady (2009). Leptanilloides chihuahuaensis differs from each of these by lacking submarginal cells and a stigma in the forewing. Geographically, L. chihuahuaensis is the only species thus far know to occur in the United States. Description (male): Body size minute. Color light yellowish-brown, head and margins of abdominal segments IV���VII darker, appendages (antennae, mandibles, legs) lighter. Integument mostly smooth and shiny, with simple curved to strongly curved suberect to decumbent setae; pilosity on head, mesosoma, petiole, and first gastral tergite (AIII) scattered and not obscuring the shininess of the integument; pilosity denser on remainder of gaster and appendages. Head in full-face view excluding eyes about as long as wide, widest above eyes; posterior corners of head evenly rounded. Eyes large, bulging, occupying almost half the side of head. Ocelli small, protruding slightly, arranged in almost an equilateral triangle, but with distance between lateral ocelli slightly greater than distance between lateral ocellus and median ocellus; distance between median ocellus and eye approximately the length of eye. Clypeus short, tranverse, lacking lamelli; lateroclypeal teeth and hypostomal teeth lacking. Mandible slender, falcate; apex blunt; masticatory margin edentate; external margin of mandible mostly evenly curved along its length; mandible tips overlap at closure. Antennal sockets circular and exposed, located at the anterior clypeal margin with anterior edge of sockets slightly overlapping anterior clypeal margin. Antenna 13 -merous; scape, pedicel, and each flagellomere longer than wide. Scape length about twice the length of the first flagellomere, and about the combined length of the first and second flagellomere; scape subequal to the length of ultimate flagellomere; pedicel thickened, length about 1.5 times width; flagellomeres 1-11 each at least twice as long as wide; first and second flagellomeres subequal in length. Pronotum U-shaped in dorsal view, reduced anteromedially to a thin horizontal strip, set below the level of the dorsally protruding mesonotum; pronotum triangular in lateral view, with pointed posterior apex directed toward wing base. Mesoscutum lacking notauli; parapsidal lines not discernable. Axillae depressed, not meeting medially, connected by a narrow furrow. Tegula, or tegula-like structure present at wing base, minute, inconspicuous, with 2- 3 erect setae. Mesopleuron lacking oblique transverse sulcus, not divided into anepisternum and katepisternum. Mesoscutellum prominently bulging, as seen in lateral view. Metapleural gland not discernable. Propodeum with dorsal and declivious surfaces not differentiated, evenly rounded. Propodeal spiracle small, ovate, positioned slightly below midheight of propodeum and slightly posterior to the midlength. Legs slender, mesotibia and metatibia each with two simple spurs, pretarsal claw lacking preapical tooth. Wings with reduced venation; clear; fringed with short to longer fine setae; with numerous short, fine microsetae evenly distributed across wing surfaces and with scattered longer setae present. Fore wing: Costal (C) vein present, about �� the length of wing, tubular basally for less than �� the length of wing, then becoming nebulous before fading out completely. Pterostigma not present. Subcostal and radial veins fused forming Sc+R; located just below and parallel to costal vein, approximately the same length as costal vein and similarly tubular for about �� of its length before becoming nebulous. Medial (M) and cubital (Cu) veins fused (M+Cu), nebulous, slightly less than �� of the wing length. Anal (A) vein present beneath M+Cu, about the same length as M+Cu, tubular for slightly more than half of its length before becoming nebulous; apically M+Cu connected to A by nebulous crossvein cu-a; subbasal cell present between M+Cu and A. Branching upward from the top of Cu-a is the tubular abscissa Rs+M (the joined veins M��f 1 and Rs+M). Radial sector (Rs), tubular, branches upward from Rs+M, then becomes nebulous just past the nebulous/spectral crossvein 2 r-rs (1 r-rs is absent). M branches off of Rs+M just before Rs, appearing spectral basally then continuing as a nebulous vein that does not reach the wing margin. Cu is present as a nebulous vein that branches off near the apex of A; Cu does not reach the wing margin. Posterior margin of fore wing with narrow, conspicuous fold where hamuli attach. Hind wing with Sc+R present, tubular basally before becoming nebulous, less than �� the length of wing. No other veins present. Anterior margin of hind wing past midlength with a thin, but conspicuous dark stigma. Three hamuli present in the region of the stigma. Jugal lobe absent. Petiole (abdominal segment II) subquadrate in lateral view, about as long as high or wide, not constricted posteriorly; in dorsal view anterior corners widened, sides slightly constricted, appearing shallowly concave before widening posteriorly; petiole broadly joined to abdominal segment III; petiolar spiracle located on anterior third of the segment, near anterodorsal extremity. Abdominal segment III larger than petiole, not developed as postpetiole, and not separated from abdominal segment IV by a constriction. Abdominal spiracles III, IV, V, and VI located on anterior third of tergites (abdominal spiracle VII not visible in examined specimens). Abdominal tergite VIII (pygidium) small, simple and visible dorsally, not entirely covered by abdominal tergite VII. Pygostyli absent. Abdominal sternite IX (subgenital plate) with posterior margin broadly concave, but with a median subtriangular process. Paramere (basimere + telomere) small (about the length of petiole); basimere roughly triangular, widest at base, narrowing to incurved falcate telomere that terminates in a blunt apex. Volsella triangular, with slightly rounded apex; volsella not differentiated into digitus and cuspis. Penisvalva elongate triangular, extending to about as far as paramere apex. Species Country GenBank# BOLD # COI Size (bp) p-distance (%) Analysis of COI. A 658 bp region of the mitochondrial gene COI, the barcode region, was sequenced successfully and the resulting consensus sequence was submitted to GenBank (Table 1). Five additional COI sequences for Leptanilloides were found and downloaded from the BOLD database (Table 1), including three for L. gracilis Borowiec & Longino (southern Mexico, Honduras), one for L. nubecula Donoso et al. (Ecuador), and one for an undescribed species L. MAS001 (Costa Rica). Comparing L. chihuahuaensis to the other specimens, pairwise genetic distances were all above 8.0%, ranging from 8.2 % (L. gracilis, Honduras) to 20.1 % (L. nubecula, Ecuador)., Published as part of Macgown, Joe A., Schiefer, Terence L. & Branstetter, Michael G., 2015, FIrst record of the genus Leptanilloides (Hymenoptera: FormIcIdae: DorylInae) from the UnIted States, pp. 392-400 in Zootaxa 4006 (2) on pages 394-398, DOI: 10.11646/zootaxa.4006.2.10, http://zenodo.org/record/58997, {"references":["Kusnezov, N. (1953) Las hormigas en los Parques Nacionales de la Patagonia y los problemas relacionados. Anales del Museo Nahuel Huapi, 3, 105 - 124.","Longino, J. T. (2003) A new Costa Rican species of Leptanilloides (Hymenoptera: Formicidae: doryline section: Leptanilloidinae). Zootaxa, 264, 1 - 6.","Donoso, D. A., Vieira, J. M. & Wild, A. L. (2006) Three new species of Leptanilloides Mann from Andean Ecuador (Formicidae: Leptanilloidinae). Zootaxa, 1201, 47 - 62.","Ward, P. S. & Brady, S. G. (2009) Rediscovery of the ant genus Amyrmex Kusnezov (Hymenoptera: Formicidae) and its transfer from Dolichoderinae to Leptanilloidinae. Zootaxa, 2063, 46 - 54."]}

Massonia amoena Mart. -Azorín, M.Pinter & Wetschnig, sp. nov. (Figs. 2–9) Species notabilis combinatione unica characterum ab omnibus speciebus Massoniae differt foliis glauco-viridulis, supra numerosas pustulas viridulas, purpurascentes atque rubescentes obsitis, quae valde heterogeneis sunt (aliquas minutas et circulares, ceteras oblongas et longitudinaliter dispositas); segmentis perigonii per anthesin valde reflexis a basi vix sigmoideis; filamentis in tubi breve 1−2.5 mm supra perigonium connatis; antheris et polline omnibus cyanellis; atque ovario in stylo gradualiter desinente. Type:— SOUTH AFRICA. Eastern Cape: Northern slopes Andriesberg, Alt. 5500 ft. [approx. 1680 m], May 1899, Fl. White, E. E. Galpin 2612 (holotype GRA!, Fig. 2). Herbaceous perennial bulbous geophyte. Roots branched, usually present for about two vegetation periods. Bulb ovoid, ca. 15−20 × 10−15 mm, inner scales fleshy and white, outer tunics papery and brownish. Leaves 2, deciduous, leaf blades opposite, spreading and appressed to the ground, 2−8 × 1−5 cm, synanthous, ovate with acute apex, with a short apicule ca. 1 mm long, with minutely papillose or shortly ciliate margin, narrowed into a subterraneous petiole 1−3 cm long that clasp the inflorescence and the peduncle; adaxial side glaucous green with numerous (50−150 per cm 2), dark green, purplish or reddish emergences, being heterogeneous in morphology, from circular (0.3−0.8 mm diameter) to elongated and disposed longitudinally (1−3 × 0.4−0.8 mm), 0.2−0.4 mm high with several short papillae on top; abaxial side green, smooth. Inflorescence a dense, subcapitate raceme, up to 1−2 cm long, with 8−32 flowers, shortly overtopping ground level, but flowers usually long exerted. Bracts membranous, green, sometimes with a purplish flush in the upper half and white below, glabrous with entire margins; lower bracts ovate 14−21 × 10−13 mm; upper bracts narrowly ovate, 12−16 × 5–8 mm. Pedicels 4−8 mm long. Flowers proterandrous, tubular, actinomorphic, with a strong pleasant smell. Perigone white or rarely pinkish, free segments deltoid or narrowly triangular, 4–7 × 1.5–3 mm, white, first straight and erect, later spreading and finally strongly reflexed at anthesis with a very slight curve at the base. Perigone-filaments tube 15−24 × 2.5−3.5 mm at anthesis, cylindrical, white or pinkish. Filaments white or tinged with pink, free portions narrowly triangular, 1–4 mm long, rather fleshy and thickened, straight, erect, connate at the base for ca. 1−2.5 mm above the perigone to form a cylindrical, filaments tube. Anthers ca. 2 mm long when closed, oblong, with dark blue anther wall, dorsifixed. Pollen dark blue. Gynoecium cenocarpous-syncarpous, narrowly obclavate, with septal nectaries; nectar colourless, sometimes yellow when aged. Ovary oblong, green, 3.8– 4.8 × 1.8–2.2 mm, with 10–14 ovules/locule; style white, 9–17 × 1–1.4 mm, thick, narrowly triangular, erect, gradually tapering to the punctiform stigma, shorter or as long as the perigone-filaments tube. Capsule loculicidal, 10–14 × 5–8 mm, valves splitting down to the base, ovate-oblong in lateral view and trigonous in apical view. Seeds black, with a greyish overlay (epicuticular wax), somewhat glossy, 2–2.4 × 1.6–2 mm, ellipsoidal, flattened at the chalazal region, with an inclined, conical apex at the micropylar region. (Figs. 3–9). Taxonomic relationships: — Massonia amoena can be easily distinguished from all other species in the genus by the glaucous green adaxial side of the leaves with numerous, dark green, purplish or reddish, morphologically heterogeneous emergences, these being from small and circular to elongated and disposed longitudinally; the strongly reflexed perigone segments at anthesis with a slight sigmoid curve at the base; the filaments connate for 1−2.5 mm above the perigone; the blue anthers with blue pollen; and the style gradually tapering from the ovary (Figs. 3–9). Its closest known relative appears to be Massonia jasminiflora, which shares the long perigone-filaments tube, the connate filaments, the blue anthers and pollen and the pleasant sweet smell. However, M. jasminiflora differs in the smooth leaves, commonly ciliate, the patent free portions of the perigone segments at anthesis, and its allopatric distribution (Fig. 1, 10; Table 2). Massonia wittebergensis and M. saniensis share with M. amoena the leaves bearing emergences, the reflexed perigone segments, the connate filaments and blue anthers, but both species differ by the shorter perigonefilaments tube, the yellow pollen, the very different morphology of leaf emergences and their allopatric distribution (Table 2, Fig. 10). Etymology:—Species named after the striking and beautiful leaves and flowers (amoenus,– a,– um: pleasant, delightful). Biology: — Massonia amoena starts to emerge from dormancy during late summer in its natural habitat. Plants grow mostly from late February to early April and by the end of April (autumn in the southern hemisphere) leaves have reached their maximum development. Flower buds begin to form as early as February but they do not start to develop properly until April (Craib & Knoll 2000). Massonia amoena flowers mainly in May and June in the wild. In cultivation in Europe they flower from late October to December. Craib & Knoll (2000) commented that honey bees and a variety of flies were observed visiting flowers of cultivated plants, suggesting that there may be more than one pollinating agent in wild populations. The very intense pleasant smell, the long perigone-filaments tube and the white flower colour would suggest pollination by moths. A more detailed study based on wild populations is necessary to evaluate these statements. Habitat: —The new species occurs at elevations from approximately 1300 to 2300 m, mainly in seasonally moist seepage areas facing south, west or east. It is usually found on rocky slopes at the base of large sheets of exposed dolerites, basalt or sandstone, but also in flat open grassveld with rocky patches. These habitats experience cold weather with severe frost in May and June at the peak of Massonia amoena flowering season. Temperatures are often well below freezing and snow may fall on the mountains (Craib & Knoll 2000). All known populations of Massonia amoena are confined to the Grassland Biome sensu Mucina & Rutherford=2006(). The northern populations from the Witteberge and the Stormberg areas are included in the Drakensberg Grassland vegetation unit. The populations from the Witteberge are located in the Southern Drakensberg Highland Grassland (Gd4) and Lesotho Highland Basalt Grassland (Gd8), whilst the Stormberg population occurs in the Stormberg Plateau Grassland (Gd3). The central populations from the Toorberg, Andriesberg and Bowker’s Kop are included in the Sub-Escarpment Grassland vegetation unit, and the Queenstown Thornveld (Gs16) and the Tarkastad Montane Shrubland (Gs17) sensu Mucina & Rutherford=2006(). The populations from the Great Winterberg are located in the Dry Highveld Grassland vegetation unit, and the Karoo Escarpment Grassland (Gh1). Finally, the southernmost population in the Amathole Mountains occurs in the Drakensberg Grassland vegetation unit and the Amatole Mistbelt Grassland (Gd2). These regions show bimodal spring-autumn or summer rainfall, usually with very dry winters, with precipitation ranging from 500–1300 mm, and an approximate mean of 50 days with frost per year (cf. Mucina & Rutherford 2006, Clark et al. 2014). Distribution: —Known from several localities restricted to the Eastern Cape province of South Africa, from the Witteberge in the Southern Drakensberg near Lady Grey and Motkop in the north to the Great Winterberg–Amathole Ranges in the south around Cathcart and Tarkastad areas, being apparently restricted between the meridians 26− 28º E (Fig. 10). Therefore, the distribution of the new species can be included in the Drakensberg Alpine Centre of Endemism as redefined by Mucina & Rutherford=2006(). The close related Massonia jasminiflora shows an allopatric distribution (Fig. 10), not known to us to occur in the Eastern Cape province, although some of their populations are located near the border to the latter province. The main distribution of M. jasminiflora is located in central and southern Free State province, with some populations extending to the adjacent areas in the North West province and Northern Cape province. It is worth mentioning that one apparently disjunct population of M. jasminiflora occurs in Mpumalanga (Fig. 10), near the golf course in Middelburg (Nieuwoudt 273 PRE), although the quarter degree given in the herbarium label (3125AC) seems to be mistaken with Middelburg from the Eastern Cape province. On this respect, Retief & Herman (1997) already cited M. jasminiflora (van Wyk coll. nº 928) from an area between Middelburg and the Waterberg in northeastern South Africa. Morphological variation: —A certain variation on leaf morphology in Massonia amoena (under the name M. jasminiflora) has been reported by Craib & Knoll (2000), especially regarding pustule size, colour and morphology. Craib & Knoll (2000) described the population at the type locality (Andriesberg) having always pustulate leaves, however some leaves were described as tinged with mauve or reddish brown and the pustules being ruby red, brown or mauvish brown. On the contrary, Craib & Knoll (2000) described the population from the Toorberg near Tarkastad as having nearly all plants pustulate leaves, but a few were plain with the surface merely tinged with various shades of mauve and sienna brown. Among the pustulate-leaved plants, specimens existed with big, evenly spread, mauvish pustules, some of them with congested ruby red pustules as well as other plants so densely pustulate that the leaf surfaces resembled coarse gravel sandpaper. It is also worth mentioning that the collection Galpin 1817 (K000257151!) from Bowker’s Kop, Queenstown, which is mounted together with the type of M. jasminiflora, includes four flowering plants that agree in flower morphology with M. amoena, but some of the pressed leaves apparently do not show evident pustules. However, a closer investigation of this collection reveals that the specimen situated at the bottom was mounted with the leaves showing the abaxial side and at least the plant placed on the left side show several small pustules on the leaves. Furthermore, a duplicate of Galpin 1817 (PRE0050981-0) includes 12 plants with strongly reflexed perianth segments and leaves showing evident remnants of pustules. In this regard, pressed leaves of Massonia amoena sometimes do not retain the morphology of the raised pustules, but remnants of them remain and the short papillae on top of each are visible. Craib & Knoll (2000) proposed that “the pustulate-leaved form, which is particularly cryptic, has probably evolved to mimic gritty dolerite soil. Evidence of this was found in the Toorberg plants, where very few had plain leaves and these were also either tinged with mauve or brown, causing them to blend better with the colours of the microhabitat”. As a summary, all living specimens of M. amoena examined in this study showed evident pustules, being constantly heterogeneous in morphology from small and circular to elongated longitudinally, where a slight variation on pustule and leaf colour was detected (Fig. 3). However, in general terms, the leaf morphology of M. amoena can be considered as constant, especially regarding the diagnostic character differing from M. jasminiflora (Fig. 1, 3). We were not able to study living specimens of Massonia amoena with smooth leaves, as cited by Craib & Knoll (2000), but we suggest that these forms could be the outcome of possible hybridization processes with other species of Massonia bearing smooth leaves, such as for instance Massonia huttoni Baker (1870: 390), M. versicolor Baker (1876: 184) and M. modesta Fourcade (1932: 79), which also occur in the Eastern Cape Province of South Africa, and which are sympatric at least with some of the southernmost populations of M. amoena. However, a detailed study on this aggregate of taxa is necessary to evaluate this. Additional specimens studied (paratypes): — SOUTH AFRICA. Eastern Cape. Lady Grey (3027 CB): Lady Grey, Joubert’s Pass, Alt. 2237 m, 20 February 2011, WW04486 (GZU!); Lady Grey (3027 CD): Motkop, Alt. 2044 m, 20 February 2011, WW04485 (GZU!); Lady Grey (3027): Majuba Nek, Herschel Distr., January 1916, Hepburn 271 (GRA!); Queenstown (3126 BA): Stormberg area (between Molteno and Dordrecht), Farm Streep Fontein 237, Alt. 1961 m, open rocky area on summit of ridge, growing among basalt boulders, rare, E. L . Gaisford & V. R. Clark 333 (GRA!); Queenstown (3126 CD): in the foothills of the Toorberg, near Tarkastad, C. Craib & C. Knoll (Photo!); Queenstown (3126 DA): Andriesberg, Alt. 1777 m, 19 February 2011, WW04484 (GZU!); Queenstown (3126 DA): Northern slopes Andriesberg, Alt. 5500 feet, May 1899, Fl. white, E. E . Galpin 2612 (GRA!); Queenstown (3126 DB): Sterkstroom, Stapelbergskloof, Halseton, SW solpes of doleritic range, in cracks of rock, Alt. 6400 feet, 28 April 1963, C. R . Callaghan 32 (PRE0050982 -0!); Queenstown (3126 DD): Bowker’s Kop, Queenstown, Alt. 4000 feet, May 1894, Fl. white, E. E . Galpin 1817 (K000257151!, PRE0050981!); Lady Frere (3127 AC): Tafelberg, Dordrecht Distr., on farm Blacks Siding, 25 May 1964, R . D. Bayliss 2237 (PRE0050983 -0!); Lady Frere (3127 BC): Top of Cala Pass, 09 March 1987, Glen 1720 (PRE0727264 -0!); Fort Beaufort (3226 AB): Great Winterberg (Tarkastad District), Farm Newtondale 228, cliffs and plateau edge, aspect NE, loamy clay, stony soil/rocky, grassland, dolerite, on cliffs, Alt. 1733 m, 18 March 2011, V. R . Clark & G. Neef 79 (GRA!); Fort Beaufort (3226 AC): Along De Beers Pass, Great Winterberg (Tarkastad District), aspect S, hill slope, loam, stony soil/rocky, sandstone, rare, along the road cutting on pass, Alt. 1605 m, 17 March 2011, V. R . Clark & G. Neef 10 (GRA!); Fort Beaufort (3226 AD): Great Winterberg Mountain, flat shelf on SW-facing slope, Alt. 1885 m, 24 May 2014, T . van Niekerk s.n. (GRA!); Fort Beaufort (3226 BD); Amathole Mountains, ex. Simply Indigenous, WW 02266 (GZU!)., Published as part of Martínez-Azorín, Mario, Pinter, Michael, Deutsch, Gerfried, Brudermann, Andreas, Dold, Anthony P., Crespo, Manuel B., Pfosser, Martin & Wetschnig, Wolfgang, 2014, Massonia amoena (Asparagaceae, Scilloideae), a striking new species from the Eastern Cape, South Africa, pp. 121-137 in Phytotaxa 181 (3) on pages 126-134, DOI: 10.11646/phytotaxa.181.3.1, http://zenodo.org/record/5145939, {"references":["Craib, C. & Knoll, C. (2000) Massonia jasminiflora: A beautiful species from the climatically severe interior of South Africa. Herbertia 55: 140 - 143.","Mucina, L. & Rutherford, M. C. (Eds.) (2006) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria. 807 pp.","Clark, V. R., Dold, A. P., McMaster, C., McGregor, G., Bredenkamp, C. & Barker, N. P. (2014) Rich sister, poor cousin: Plant diversity and endemism in the Great Winterberg - Amatholes (Great Escarpment, Eastern Cape, South Africa). South African Journal of Botany 92: 159 - 174.","Retief, E. & Herman, P. P. J. (1997) Plants of the northern provinces of South Africa: keys and diagnostic characters. National Botanical Institute, Pretoria, pp 681.","Baker, J. G. (1870) A revision of the genera and species of herbaceous capsular gamophyllous Liliaceae. Journal of the Linnean Society Botany 11: 349 - 436.","Baker, J. G. (1876) On new bulbous plants from the eastern provinces of Cape Colony. Journal of Botany, British and Foreign 14: 181 - 184.","Fourcade, H. G. (1932) Contributions to the flora of the Knysna and neighbouring divisions. Transactions of the Royal Society of South Africa 21: 75 - 102."]}

Oligoryzomys brendae Massoia, 1998 Type locality: ������Tucum��n (localidad t��pica: Cerro San Javier, Dpto. Taf�� Viejo) (aprox. 1000 m. de altura)������ = Argentina, Tucum��n, Taf�� Viejo, Cerro San Javier (ca. -26.78��, -65.37��). Emended diagnosis. A member of the sigmodontine genus Oligoryzomys characterized by the following combination of characters: size large for the genus (HBL: 90 mm; CIL: 22.9, MTL: 3.9, BB: 12.1; all measurements in average); dorsal coloration orangish brown, with somewhat grizzled appearance; ventral hairs basally grey and whitish to ochraceous at the tip; ears short, rounded, and dark brown; tail longer than head and body and sharply bicolored; skull (Fig. 3) relatively robust, with a short and broad rostrum, well expanded zygomatic arches and a braincase inflated and broad; interorbital constriction narrow and hourglass shaped, with slightly defined supraorbital ridges; incisive foramina relatively long, posteriorly extended to the anterior border of M 1; karyotype with 2 n = 58, FN = 74. Distribution. Oligoryzomys brendae is mainly found in forested and highland grassland environments of the Yungas between 700 and 2900 m elevation, from northernmost Salta province southward to Catamarca, Argentina. Additionally, we recently recorded the species further south in La Rioja province where it is restricted to isolated humid ravines at the ecotone between the xeric Chaco Seco and Monte desert formations. Jayat et al. (2011 b) cited a large species of Oligoryzomys for the Famatina range in La Rioja province; nevertheless, the specimens from Pampa de la Viuda reported here (see Appendix 2) constitute the first unequivocal mention of O. brendae for the province of La Rioja. Description. Oligoryzomys brendae is a large-sized species within the genus (Figs. 3, 4; Table 1); its dorsal pelage is soft, and dense; individual hairs have gray bases and ochraceous to reddish tips, giving a general grizzled orangish-brown appearance, slightly darker at the head and midline and orangish towards the cheeks and flanks. Cover hairs are 10 to 12 mm long, being larger in the mid rump region; guard hairs are longer and darker, projecting 3���4 mm beyond the fur in the rump. Ventral hairs are basally grey and whitish to ochraceous at the tip. Ears are short, rounded, and dark brown in coloration; its inner surface is covered by short brown hairs with yellowish tips. Eyes are large and are surrounded by a conspicuous ring of dark brown hairs. Mystacial vibrissae are abundant and moderately long, reaching or slightly surpassing the base of the ear. Interamal and submental vibrissae are short and white. Fore and hindfoot are dorsally covered by short white hairs. Ungueal tuft are whitish and slightly larger than claws. The tail is longer than head and body and sharply bicolored, dark grey dorsally and whitish in the belly. The skull (Fig. 3) is relatively robust, with a short and broad rostrum, well expanded zygomatic arches and braincase inflated and broad. Nasals are slightly expanded in its distal third. The interorbital constriction is narrow and hourglass shaped, with slightly defined supraorbital ridges. Temporal and mastoid crests are scarcely developed. The zygomatic notches are wide and deep. The dorsal profile of the skull is nearly convex. Zygomatic plate is broad, with its anterior margin nearly straight and the upper root slightly slanted. Long and delicate hamular process separates a well developed postglenoid foramen from a small subsquamosal fenestra. Incisive foramina are relatively long, posteriorly extended to the anterior border of M 1. Mesopterygoid fossa is broad, with straight and slightly divergent lateral borders; its anterior margin is well behind the posterior plane defined by the M 3. Posterolateral palatal pits are rounded and large, located nearly the anterior border of the mesopterygoid fossa. Parapterygoid fossae are wide and relatively deep, with convex external borders. Auditory bullae are inflated and with relatively broad and long eustachian tubes. The associate foramina of the otic capsules and cephalic arterial patterns do not differ from the typical condition seen in other Oligoryzomys species and fully described by Carleton & Musser (1989). Both specimens lack alisphenoid strut, sphenofrontal foramen and the accompanying squamosalalisphenoid groove. The mandible is robust and comparatively deep. The masseteric crest is well developed, with their anterior end situated above the mental foramen and extending at the level or slightly beyond the anterior border of m 1. The capsular projection is strongly developed and situated below a shallow sigmoid notch. The coronoid process is comparatively broad and very short, extending slightly above the level of the condyle. The angular process is also heavily constructed and does not surpass posteriorly the level of the condyle. Upper incisors are ungrooved and opistodont, with orange enamel in its anterior surface. Molars are pentalophodont and brachyodont, with the lingual cusps slightly anterior to the labial ones; the anteromedian flexus is deep and defines prominent anterolingual and anterolabial conules; the anteroloph and parastyle are comparatively short and broad; the mesoloph and mesostyle are well developed and the posteroloph is near to metacone. The M 2 is nearly as a truncate oval and closely resembled M 1 except for the lack of a well developed procingulum. The M 3 is small and in both specimens it has a large inner enamel island. Procingulum and anterolabial cingulum on m 1 are well developed; anteromedian flexid is expressed only in young specimens. The anterolophid-metastylid and mesolophid-mesostylid are very short and the posterlophid is well developed. In the m 2 the procingulum is vestigial, the mesostylid is small and the posterolophid is well developed. The m 3 is large, reaching 50 % of the m 2, with anterolabial cingulum and ectostylid vestigial. Morphological comparisons. Our examination of the holotype and paratype of O. brendae and large series of Oligoryzomys from northwestern Argentina allow us to state that there are only one large-sized form of Oligoryzomys inhabiting forested and high altitudinal grasslands areas of northwestern Argentina. This species is Oligoryzomys brendae, which differs from the sympatric O. flavescens by its larger size, darker ears, absence of white hairs in chin and throat, broader rostrum, broader zygomatic notches and more expanded zygomatic arches. Oligoryzomys chacoensis (Myers & Carleton, 1981), that is found at lower elevations and Chacoan environments, has conspicuously posteriorly divergent interorbital edges, being intermediate in size between O. brendae and O. flavescens, and having a more strong orangish dorsal fur that contrast with the whitish underside. A small but distinctive tuft of orangish hairs are anterior to the relatively larger ears. Furthermore, hairs on chin and throat are white to the base. Compared with specimens from populations referred to O. destructor, O. brendae has longer ears (17���20 mm, vs. 12���14 mm [Peru], 14���17 [Ecuador] and 15���17 [Bolivia]), larger hindfeet (24���28 mm, vs. 22.5���24.5 [Peru], 22���25 [Ecuador] and 19���25 [Bolivia]) and a wider braincase (11.5���12.6 mm, vs. 11.4 mm [Peru], 10.4���11.2 [Ecuador] and 10.4���11.8 [Bolivia]; see Osgood, 1914; Hershkovitz, 1940; Olds & Anderson, 1987; Anderson, 1997). Bolivian populations have more grayish venters (cf. Olds & Anderson, 1987), while Peruvian and Ecuatorian forms (including true O. destructor, and the nominal forms stolzmanni, spodiurus and melanostoma) are more reddish to ochraceous dorsally and brighter colored (Thomas, 1926 a, 1926 b; Hershkovitz, 1944; Olds & Anderson, 1987). It is worth noting that specimens referred by Hanson et al. (2011) to O. destructor (TEL 1479, 1481; assessed via photographs), and included in our phylogenetic analysis, match the description of the Ecuadorian populations made by Hershkovitz (1944) under Oryzomys (Oligoryzomys) spodiurus (= O. destructor, see Musser & Carleton, 2005) in having narrow nasals, expanded palatine foramina, long molar toothrows, wide palates and mesopterygoid fossae, narrow zygomatic arches and grayish tails. Genetic variation. Our sampling of O. brendae was not large enough to constitute a phylogeographic analysis, but our results provide a preliminary view on the geographic structure of the genetic variation of this species. We analyzed 12 sequences gathered at six localities from the Yungas of Catamarca, Tucum��n, Salta and Jujuy provinces and two from Monte-Chaco Seco ecotone of La Rioja province, in northwestern Argentina. These haplotypes are quite similar, differing on average by an observed value of only 0.9 %. In addition, despite of the large geographic coverage of the sample covering more than 700 km (ca. six latitudinal degrees), and several environmental discontinuities (e.g., large rivers, xeric habitats) among the analyzed populations, there is a lack of phylogeographic structure in the analyzed sample (Fig. 5). These two facts reinforce the notion that only one species of large-sized Oligoryzomys, O. brendae, inhabits eastern mountain ranges of nortwestern Argentina. The lack of phylogeographic structure of O. brendae resembles the pattern of A. spegazzini (Jayat et al. 2010), that has a similar distributional pattern, while contrasts with that shown by Necromys lactens (D'El��a et al. 2008) over roughly the same distributional range but mainly restricted to the high altitudinal grasslands of the Yungas. The clarification of causes behind these similitude and difference awaits further studies with larger samplings. Notwithstanding, the environmental history of the region (Iriondo, 1993; Ortiz et al., 2011 and references therein) and the ecological requirements of these species, with N. lactens inhabiting open and isolated highland habitats (Jayat & Pacheco, 2006), and O. brendae (Jayat et al., 2008, as Oligoryzomys sp.) and A. spegazzinii (Jayat et al., 2010) occurring in more continuous forested habitats, can explain, at least in part, the observed phylogeographic differences. Final considerations. Given the results of the phylogenetic analysis, a re-evaluation of northern nominal taxa, such as Hesperomys melanostoma Tschudi, 1844, Oryzomys stolzmanni Thomas, 1894, Oryzomys stolzmanni maranonicus Osgood, 1914 and Oryzomys (Oligoryzomys) spodiurus Hershkovitz, 1940 is needed to completely clarify the taxonomic scenario within large-bodied Oligoryzomys of the forested foothills of the eastern Andean region. It is also a pending task to determine the northern distributional limit of O. brendae, until now confidently established on northern Salta province, northernmost Argentina. Anderson (1997) referred many specimens coming from several localities of Bolivia (from the northern La Paz department to southern Tarija) to O. destructor, but the correct identity of these specimens remains to be resolved. In fact, given our phylogenetic results, these populations perhaps correspond to an unnamed form of this genus. Finally, by showing that a distinct species of Oligoryzomys, O. brendae, inhabits the Yungas of northwestern Argentina, this study reinforces the biodiversity value of this eco-region, which counts with two recently discovered endemic species of Sigmodontinae, Oxymycterus wayku and Phyllotis anitae (Jayat et al. 2007, 2008). Our finding emphasizes the need to intensify field work in this biome, which unfortunately is being highly modified by human activities (Brown et al. 2001)., Published as part of Teta, Pablo, Jayat, J. Pablo, Ortiz, Pablo E. & D'El��a, Guillermo, 2013, The taxonomic status of Oligoryzomys brendae Massoia, 1998 (Rodentia, Cricetidae), with comments on the availability of this name, pp. 433-447 in Zootaxa 3641 (4) on pages 440-444, DOI: 10.11646/zootaxa.3641.4.9, http://zenodo.org/record/222692, {"references":["Massoia, E. (1998) Roedores vinculados con virosis humanas en la Republica Argentina. Capitulo 6, Vectores y Reservorios. In: (2 do. Congreso Argentino de Zoonosis, 1 er Congreso Argentino y Latinoamericano de Enfermedades Emergentes y Asociacion argentina de Zoonosis, eds.) Temas de Zoonosis y Enfermedades Emergentes. Buenos Aires, Sociedad Argentina de Zoonosis, pp. 243 - 246.","Carleton, M. D. & Musser, G. G. (1989) Systematic studies of oryzomyine rodents (Muridae, Sigmodontinae): A synopsis of Microryzomys. Bulletin of the American Museum of Natural History, 191, 1 - 83.","Myers, P. & Carleton, M. D. (1981) The species of Oryzomys (Oligoryzomys) in Paraguay and the identity of Azara´s \" Rat sixieme ou rat a tarse noir \". Miscellaneous Publication of the Museum of Zoology, University of Michigan, 161, 1 - 41.","Hershkovitz, P. (1940) Four new oryzomyine rodents from Ecuador. Journal of Mammalogy, 21, 78 - 84.","Anderson, S. (1997) Mammals of Bolivia, taxonomy and distribution. Bulletin of the American Museum of Natural History, 231, 1 - 652.","Hanson, J. D., Utrera, A. & Fulhorst, C. F. (2011) The delicate pygmy rice rat (Oligoryzomys delicatus) is the principal host of Maporal virus (family Bunyaviridae, genus Hantavirus). Vector Borne Zoonotic Diseases, 11, 691 - 696.","Musser, G. M. & Carleton, M. D. (2005) Superfamily Muroidea. In: Wilson, D. E. & Reeder, D. M. (Eds.). Mammal species of the world: A taxonomic and geographic reference Third ed. Johns Hopkins University Press, Baltimore, pp. 894 - 1531.","Iriondo, M. H. (1993) Cambios climaticos en el noroeste durante los ultimos 15.000 anos. In: Iriondo, M. (ed.). El Holoceno en la Argentina. Cadincua, Buenos Aires, pp. 35 - 44.","Jayat, J. P. & Pacheco, S. E. (2006) Distribucion de Necromys lactens y Phyllotis osilae (Rodentia: Cricetidae: Sigmodontinae) en el noroeste argentino: modelos predictivos basados en el concepto de nicho ecologico. Mastozoologia Neotropical, 13, 69 - 88.","Jayat, J. P., D'Elia, G., Pardinas, U. F. J. & Namen, J. G. (2007) A new species of Phyllotis (Rodentia, Cricetidae, Sigmodontine) from the upper montane forest of the Yungas of northwestern Argentina. In: Kelt, D., Lessa, E. & Salazar-Bravo, J. (Eds.), The Quintessential Naturalist: Honoring the Life and Legacy of Oliver P. Pearson. California: University of California Publications in Zoology, pp. 775 - 798.","Brown, A. D., Grau, H. R., Malizia, L. R. & Grau, A. (2001) Argentina. In: Kappelle, M. & Brown, A. D. (Eds.), Bosques Nublados del Neotropico. Costa Rica: Instituto Nacional de Biodiversidad, pp. 623 - 659."]}

Akodon caenosus Thomas, 1918 Akodon diminutus Barquez, D��az and Goytia, 1994. Nomen nudum (see Galliari et al., 1996). Akodon aliquantulus D��az, Barquez, Braun and Mares, 1999. Journal of Mammalogy, 80:788. Holotype: BMNH 18.1.1.38, adult male. Type locality: Le��n, Jujuy, 1500 m (Thomas, 1918). Description: Detailed morphological description in Myers et al. (1990) (as A. puer caenosus). Additional morphological and morphometric data for some populations of northwestern Argentina can be found in Thomas (1918; original description of A. puer caenosus), Barquez et al. (1980) and D��az (1999). Here we summarize the characteristics of populations along the latitudinal and altitudinal gradients of northwestern Argentina. Akodon caenosus is the smallest species of the boliviensis group in northwestern Argentina. Dorsal coloration uniform and highly variable, ochraceous brown with yellowish, rufous, or olivaceous casts. Ears of the same color than the dorsum and with a tuft of hairs in front. Laterals clearer than dorsum and more richly colored in some specimens. Venter paler, whitish gray, buffy gray, yellowish or even ruddy, clearly contrasting with dorsum. Chin with few white hairs that do not form a conspicuous patch. Inguinal region with a more intense tinge than the rest of the venter in some specimens. Fore and hind feet covered with whitish or buffy hairs. Tail strongly bicolored, dorsally blackish-brown and ventrally whitish or buffy, densely or sparsely covered depending on the analyzed populations and individuals. The skull is the smallest among the Akodon species of northwestern Argentina. Rostrum short, with very narrow and shallow zygomatic notches and frontal sinuses not well developed. Interorbital region comparatively broad, with hourglass shaped and with margins rounded or slightly squared. Zygomatic arches not flared, braincase small and inflated, with temporal and lambdoid crests not well developed. Zygomatic plate narrow with anterior border straight or slightly concave and generally slopes gently backward from bottom to top. In most of the studied specimens the posterior ascending process of alisphenoid does not reach the squamoso-alisphenoid groove or only touch their lower margin. Hamular process of squamosal delicate, but in a few individuals it is strongly built. Incisive foramina generally extended to the protocone of M1, but in some individuals its reach the hypoflexus or even further back. Mesopterygoid fossa very narrow, its anterior margin rounded or slightly squared and with lateral margins straight and slightly divergent backward. Parapterygoid fossa generally broader than mesopterygoid, with convex border and slightly divergent backward. Mandibular ramus very delicate. Masseteric crest extends slightly behind the anterior margin of m1. Capsular projection generally poorly developed and situated at the same level or behind the coronoid process. Angular process is less extended backward than the condyle. Upper incisors orthodont or slightly opistodont. The procingulum and anteromedian flexus of M1 are well developed. Anteroloph and mesoloph are always present and in some individuals an enteroloph can be observed. The M2 has a conspicuous mesoloph and the M3 shows great variability. In old individuals it is completely oval shaped but in young ones anteroflexus, metaflexus and hipoflexus are present. The m1 has the procingulum and anteromedian flexid well developed and presents protostylid and ectostylid. In some individuals a remnant of metastylid and mesostylid can be observed. The m2 retains only the labial stylid although some specimens have a very small ectostylid. The m3 presents deep labial and lingual flexids. Some individuals show a small protostylid. Karyotype: 2n = 34, FN = 40, based on six specimens from Le��n, Jujuy, and two from El Cadillal, Tucum��n (Barquez et al., 1980; Vitullo et al., 1986 as A. puer). Variation: We have observed substantial morphological variation for this species. Within the same population, we find pale and dark individuals, some of them with striking rufous tones, particularly lactating females. Among the cranial characters, the development of zygomatic notches, zygomatic plate, and the thickness of hamular process show some variability. We also observed morphometric variations for individuals of the same age and population, with remarkable differences among some of the studied localities. In addition, consistent variation in the coloration pattern with elevation and some geographic variation for some morphometric variables are also apparent. The specimens from high altitude populations are conspicuously paler than those from lower localitites, showing more buffy tinges in the coloration of the fur and more developed eyerings. These populations come from open environments characterized as the ecotone between humid grasslands and semiarid areas (e.g., the Prepuna or impoverished grasslands of high-Andean region). Specimens from central Salta and central Tucum��n provinces are on average smaller than individuals from B��rcena and Reyes, southern Jujuy, near the type locality. Comparisons: Externally this species can be confused with A. boliviensis and young individuals of A. spegazzinii. The differences between A. caenosus and A. boliviensis were described in detail by Myers et al (1990) and are summarized above. In areas of sympatry A. caenosus and A. spegazzinii follow similar variation patterns in coloration through altitudinal gradients. However, many morphometric values and several cranial characters are useful in specimen determination. In 15 of the 20 analyzed morphometric variables, A. caenosus is significantly smaller than A. spegazzinii (Tables 1 and 2). This is also verified by the minimum overlap in the PCA analysis (Fig 2) and the lack of missclasifications in the DA. In addition, Akodon caenosus shows narrower and shallower zygomatic notches, comparatively broader interorbital constriction, temporal and mastoid ridges less developed, narrower mesopterygoid fossa and less hypsodont molars. The average percentage of genetic divergence between these species is the greatest (7.7%) observed for the boliviensis group in northwestern Argentina (Table 12). Akodon caenosus and A. sylvanus represent two extremes in the morphometric variation observed within the boliviensis group in northwestern Argentina (Tables 1 and 2; figure 2). Akodon caenosus is, furthermore, paler than A. sylvanus, showing a more evident contrast between dorsum and venter, and has more developed eyerings. The skull of A. caenosus has a shorter and narrower rostrum, less swollen frontal sinuses, narrower braincase and interorbital region and a narrower mesopterygoid and parapterygoid fossa. The upper incisors in A. caenosus are less orthodont and the molars less hypsodont than A. sylvanus. Additionally, these species show a high average percentage of genetic divergence (6.7%). The differences between A. caenosus and the new species are listed under the treatment of the latter. Distribution: Akodon caenosus is the more broadly distributed species of the boliviensis group in northwestern Argentina, with records from northernmost Salta to southern Catamarca, from 400 m to 3100 m elevation (Fig. 7). Habitat: Most of the records come from Yungas environments, from the lower altitudinal belt to high altitude grasslands. Notwithstanding, we have recorded the species in Chacoan environments near the ecotone with Yungas and in the lower altitudinal limit of the High Andean grasslands. Natural history: Although we have recorded specimens in reproductive condition throughout the year, most of the individuals were active between November and January. We observed the highest proportion of molting animals in fall and winter. In their broad distributional range A. caenosus can be found coexisting with many different species depending on the altitudinal or latitudinal sector in northwestern Argentina. Thus, in the north A. caenosus was captured alongside Akodon sylvanus, A. simulator, Oxymycterus paramensis, Oligoryzomys cf. O. flavescens and Oligoryzomys sp. in high altitudinal grasslands at 1400 m. At the same latitude in upper belts above 2000 m, A. caenosus is sympatric with A. boliviensis, Necromys lactens, N. amoenus, Calomys musculinus (Thomas), Phyllotis caprinus Pearson, P. osilae and P. xanthopygus. In the south, in Yungas areas of Tucum��n province, the species was caught together with Akodon spegazzinii, A. simulator, Necromys lactens, N. lasiurus (Lund), Oxymycterus paramensis, Oxymycterus wayku Jayat et al., Oligoryzomys cf. O. flavescens, Oligoryzomys sp., Phyllotis anitae, P. osilae, Andinomys edax and Abrothrix illutea. At this same latitude in Chacoan environments in transition with Yungas A. caenosus is sympatric with Akodon spegazzinii, A. simulator, Necromys sp., Oligoryzomys cf. O. flavescens, Oligoryzomys sp., Calomys sp. and Graomys centralis. Comments: Although originally described as a subspecies of Akodon puer (Thomas 1918), the form caenosus was soon considered as a valid species by Thomas (1920); this latter suggestion was followed by Cabrera (1961), Barquez et al. (1980), and Mares et al. (1981). Vitullo et al. (1986) and Apfelbaum & Reig (1989) considered specimens from Le��n, Jujuy Province (type locality of caenosus) as A. puer and Myers et al. (1990) followed this position (but see Hershkovitz 1990), considering caenosus as the subspecies of A. puer inhabiting northwestern Argentina (see also Barros et al. 1990). Anderson (1997) also recognized three subspecies for A. puer but noted the priority of the name lutescens over puer. Since then, the nomenclature used for this species has been constantly changing. Capllonch et al. (1997) and Mares et al. (1997) listed caenosus as valid species whereas D��az et al. (1999, 2000) recognized caenosus as subspecies of puer for the populations of the northwestern Argentina. Notwithstanding, D��az (1999) and D��az & Barquez (2007) considered again the nominal form caenosus as representing a valid biological species including A. lutescens puer as an additional sigmodontine species in northwestern Argentina. Musser & Carleton (2005) and Pardi��as et al. (2006) followed the conclusions of Myers et al. (1990). D��az et al. (1999) described Akodon aliquantulus, from the ecotone between upper Yungas forest and highland grassland in Tucum��n province (Las Ag��itas, Cumbres del Taficillo, 1700 m, 26�� 42��� S, 65�� 22��� W), and related it to the A. boliviensis group on the basis of its morphology. The unimpressive morphological distinction from A. lutescens, based only on multivariate ordinations and univariate overlap, induced some authors to recommend a revision of the status of this form (Musser & Carleton 2005; Jayat et al., 2008a). This species was diagnosed and differentiated from A. caenosus (as A. puer caenosus) and A. spegazzinii on the account of being smaller in a number of measurements and the ���weakly developed eye ring��� of two individuals originally preserved in fluid and then prepared as skin plus skull. We could not distinguish the type specimens of A. aliquantulus from A. caenosus following the diagnosis provided by D��az et al. (1999). Most of the characters used for the diagnosis of A. aliquantulus (centered, as said, on a few external and cranial dimensions) are subtle, highly variable in many sigmodontine species, and what is most important are age-dependent. Although D��az et al. (1999) considered the two specimens as old individuals (age-class 5) our observations indicate that they must be assigned to the age class 4 of Myers (1989) (Figure 8). The morphometric distinction between A. aliquantulus and A. caenosus was based on a PCA analysis with a relatively low explained variance (63.3%, 7.9% and 5.1% for components 1, 2 and 3 respectively) and the statistic significance of the observed differences was not provided by the authors. We think that the erroneous assignment of age for the two specimens by D��az et al. (1999) misinform their conclusions. The analysis of table 1 of D��az et al. (1999) clearly indicates the overlap between A. aliquantulus specimens and A. caenosus exemplars of ageclass 4 (see also Table 9 of this work). Remarkably, even some of the diagnostic skull measurements (e.g. braincase breadth, maxillary toothrow length) are included in the observed range for age-class 5 individuals of A. caenosus. Moreover, all the characters mentioned in the description of A. aliquantulus are within the levels of morphological variation observed in A. caenosus of Northwestern Argentina. Therefore, we find no defensible argument to consider A. aliquantulus as a distinct species and consider it a junior synonym of A. caenosus., Published as part of J. Pablo Jayat, Pablo E. Ortiz, Jorge Salazar-Bravo, Ulyses F. J. Pardi��as & Guillermo D'El��a, 2010, The Akodon boliviensis species group (Rodentia: Cricetidae: Sigmodontinae) in Argentina: species limits and distribution, with the description of a new entity, pp. 1-61 in Zootaxa 2409 on pages 23-25, DOI: 10.5281/zenodo.293461, {"references":["Thomas, O. (1918) On small mammals form Salta and Jujuy collected by Mr. E. Budin. Annals and Magazine of Natural History, (9) 1, 186 - 193.","Galliari, C. A., Pardinas, U. F. J. & Goin, F. J. (1996) Lista comentada de los mamiferos argentinos. Mastozoologia Neotropical, 3, 39 - 61.","Diaz, M. M., Barquez, R. M., Braun, J. K. & Mares, M. A. (1999) A new species of Akodon (Muridae: Sigmodontinae) from Northwestern Argentina. Journal of Mammalogy, 80, 786 - 798.","Myers, P., Patton, J. L. & Smith, M. F. (1990) A review of the boliviensis group of Akodon (Muridae: Sigmodontinae) with emphasis on Peru and Bolivia. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 177, 1 - 89.","Barquez, R. M., Williams, D. F., Mares, M. A. & Genoways, H. H. (1980) Karyology and morphometrics of three species of Akodon (Mammalia: Muridae) from northwestern Argentina. Annals of Carnegie Museum, 49, 379 - 403.","Diaz, M. M. (1999) Mamiferos de la Provincia de Jujuy: Sistematica, distribucion y ecologia. Unpublished D. Phil. Thesis, Universidad Nacional de Tucuman, Argentina, 640 pp.","Vitullo, A. D., Merani, M. S., Reig, O. A., Kajon, A. E., Scaglia, O., Espinosa, M. B. & Perez-Zapata, A. (1986) Cytogenetics of South American Akodont Rodents (Cricetidae): new kariotypes and chromosomal banding patterns of argentinian and uruguayan forms. Journal of Mammalogy, 67, 69 - 80.","Thomas, O. (1920 a) A further collection of mammals from Jujuy. Annals and Magazine of Natural History, (9) 5, 195 - 196.","Cabrera, A. (1961) Catalogo de los mamiferos de America del Sur. Parte II. Revista del Museo Argentino de Ciencias Naturales \" Bernardino Rivadavia \", Zoologia, 4 (2), 309 - 732.","Mares, M. A, Ojeda, R. A. & Kosco, M. P. (1981) Observation on the distribution and ecology of the mammals of Salta Province, Argentina. Annals of Carnegie Museum, 50, 151 - 206.","Apfelbaum, L. I. & Reig O. A. (1989) Allozyme genetic distances and evolutionary relationships in species of akodontine rodents (Cricetidae: Sigmodontinae). Biological Journal of the Linnean Society, 38, 257 - 280.","Hershkovitz, P. (1990) Mice of the Akodon boliviensis size class (Sigmodontinae, Cricetidae), with the description of two species from Brazil. Fieldiana, Zoology, new series, 57, 1 - 35.","Barros, M. A., Liascovich, R. C., Gonzalez, L., Lizarralde, M. S. & Reig, O. A. (1990) Banding pattern comparison between Akodon iniscatus, and Akodon puer (Rodentia, Cricetidae). Zeitschrift fur Saugetierkunde, 55, 115 - 127.","Anderson, S. (1997) Mammals of Bolivia, taxonomy and distribution. Bulletin of the American Museum of Natural History, 231, 1 - 652.","Capllonch, P., Autino, A., Diaz, M. M., Barquez, R. M. & Goytia, M. (1997) Los mamiferos del Parque Biologico Sierra de San Javier, Tucuman, Argentina: observaciones sobre su sistematica y distribucion. Mastozoologia Neotropical, 4, 49 - 71.","Diaz, M. M., Braun, J. K., Mares, M. A. & Barquez, R. M. (1997) Key of Mammals of Salta Province, Argentina. Occasional Papers of the Oklahoma Museum of Natural History, 2, 1 - 10.","Diaz, M. M., Braun, J. K., Mares, M. A. & Barquez, R. M. (2000) An update of the taxonomy, systematics, and distribution of the mammals of Salta province, Argentina. Occasional Papers of the Oklahoma Museum of Natural History, 10, 1 - 52.","Diaz, M. M. & Barquez, R. M. (2007) The Wild Mammals of Jujuy Province, Argentina: Systematics and Distribution. In: Kelt, D. A., Lessa, E. P., Salazar-Bravo, J. & Patton, J. L. (Eds.), The Quintessential Naturalist: Honoring the Life and Legacy of Oliver P. Pearson. University of California Publications in Zoology 134, pp. 417 - 578.","Musser, G. M. & Carleton, M. D. (2005) Superfamily Muroidea .. In: Wilson, D. E. & Reeder, D. M. (Eds.) Mammal species of the world: A taxonomic and geographic reference. Third ed. Baltimore: John Hopkins University Press, pp. 894 - 1531.","Jayat, J. P., Ortiz, P. E., Teta, P., Pardinas, U. F. J. & D'Elia, G. (2006) Nuevas localidades argentinas para algunos roedores sigmodontinos (Rodentia: Cricetidae). Mastozoologia Neotropical, 13, 51 - 67.","Jayat, J. P., Ortiz, P. E. & Miotti, M. D. (2008 a) Distribucion de sigmodontinos (Rodentia: Cricetidae) en pastizales de neblina del noroeste argentino. Acta Zoologica Mexicana, 24, 137 - 177.","Myers, P. (1989) A preliminary revision of the varius group of Akodon (A. dayi, dolores, molinae, neocenus, simulator, toba and varius). In: Redford, K. H. & Eisenberg, J. F. (Eds.), Advances in Neotropical Mammalogy. Gainesville, Florida: Sandhill Crane Press, Inc., pp. 5 - 54."]}

Mantidactylus curtus (Boulenger) Series examined. ZSM 943 / 2004 (1 specimen). The single specimen was collected near Antoetra by M. Vences in the period of 18���22 January 2003 in a stream flowing through largely deforested highland grassland, with some bushes and isolated trees. The stream had a width of 1.5���3 m and included some small and shallow tributaries. Parts of the stream were of clear and fast-flowing water and rocky bottom whereas other parts, and especially the tributaries, were slow-moving, with the water largely covered by high grass tufts. Taxonomic note. Tadpoles described by Arnoult & Razarihelisoa (1967) and Blommers-Schl��sser (1979) as Mantidactylus curtus most likely belong at least partly to a different species (see section on Mantidactylus sp. aff. curtus "Ankaratra" below). The tadpole described here originates instead from a site in the general region corresponding to the type locality of M. curtus ("East Betsileo"), and its DNA sequence is close to specimens that we assign to this species (see Glaw & Vences 2007: page 237, figs. 1 a and 1 b). Description. Based on the single voucher specimen (ZSM 0943/ 2004; field number FG/MV 2003.1757; Genbank accession number EU 717879; Figure 3) in developmental stage 26, and in good state of preservation (a huge section from the dorsal tail has been excised for DNA analysis). BL 14.8 mm, TL 42.3 mm, for further measurements see Table 1. Colouration in preservative yellowish, transparent. A dark mid-dorsal yellowish line slightly visible through skin. Dark yellowish pigments concentrated dorsally besides eyes and at middle of body. Caudal fin dorsally covered to the tip with some irregular light brown patches and spots. Belly, caudal muscle and ventral fin almost uniformly transparent. In dorsal view, body shape ovoid to nearly rounded, snout elongated and flattened to tip. In lateral view, body depressed, BW 1.32 of BH, snout rounded. Nostrils elliptical and pointed to tip of snout, rimmed with a mediodorsal projection giving them a bean shape, not protuberant, positioned dorsolaterally. IND 0.38 of IOD. Nostrils, closer to tip of snout than to eyes. Eyes positioned dorsally, directed dorsolaterally, and moderately sized, ED 0.07 of BL, not protuberant. Spiracle sinistral, not visible in dorsal view, attached to body wall but its inner wall lifting up from body wall, closer to snout than to end of body, ESD 0.26 of BL. Spiracular opening oval, laterally positioned, oriented posteriorly. Intestinal spiral not visible in dorsal view, well visible in ventral and lateral views. Medial vent tube with lateral displacement, dextral. Caudal musculature rather strongly developed, upper origin well visible in dorsal view, TMH 0.76 of BH and 0.72 of MTH, fast tapering from approximately three-fourths of tail, almost reaching the tip. Dorsal fin originating at the end of first-quarter of caudal muscle, point of maximum height located after mid-tail, MTH 1.06 of BH. Margins of ventral and dextral fin slightly convex to finely rounded tail tip. Oral disc of moderate size, generalized, laterally emarginated, ODW 0.32 of BL and 0.56 of BW, transversely ovoid to nearly rounded, anteroventrally positioned, with robust dark coloured beaks, which slightly lost keratinisation. Oral disc margins not visible in dorsal view, but visible in lateral view. LTRF 5 (2���5)/ 3 (1), with additional gaps in rows A 1 and P 3, here interpreted as oral deformities. About 30 teeth per mm and a total of 115 teeth in A 2. A 5 very small, consisting of only 5 teeth on right side and 3 teeth on left of oral disc. Three small gaps in first anterior tooth row and large medial gap in second anterior tooth row. First and third posterior tooth rows with small medial gap. Upper labium with large dorsal gap in marginal papillae, upper and lower labium with two rows of papillae. Papillae with a rounded tip (some pigmented), sporadically with pointed tip (total number of papillae: 82). Upper jaw sheath M-shaped, lower jaw sheath V-shaped, both with almost equal-sized serrations., Published as part of Schmidt, Heike, Glaw, Frank, Teschke, Meike & Vences, Miguel, 2009, Description of tadpoles of five frog species in the subgenus Brygoomantis from Madagascar (Mantellidae: Mantidactylus), pp. 48-60 in Zootaxa 1988 on pages 53-54, DOI: 10.5281/zenodo.185506, {"references":["Arnoult, J. & Razarihelisoa, M. (1967) Contribution a l'etude des Batraciens de Madagascar. Le genre Mantidactylus. Adultes et formes larvaires de M. betsileanus (Blgr.), M. curtus (Blgr.) et M. alutus (Prc.). Bulletin du Museum National d`Histoire naturelle, Paris, 39, 471 - 487.","Blommers-Schlosser, R. M. A. (1979) Biosystematics of the Malagasy frogs, I. Mantellinae (Ranidae). Beaufortia, 29, 1 - 77."]}