identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03B887AC26659820FCB148F8FD9EFE75.text	03B887AC26659820FCB148F8FD9EFE75.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Otomys willani Taylor & Kearney & Dalton & Chakona & Kelly & Barker 2020	<div><p>OTOMYS WILLANI SP. NOV., WILLAN’S VLEI RAT</p><p>Holotype: Ditsong National Museum of Natural History (TM) No. 49101 (field number BAR1020) is a male, age class 3, probably young adult (sensu Taylor et al., 1993), with intact cranium (skin rotten and discarded), part of a series of specimens collected at  Tiffendell, Eastern Cape Province in December 2013 by G. Goldner. The specimen has been included in both morphometric and molecular analyses. The following external measurements were recorded: total length 198 mm, tail length 74 mm, hind length 25/ 22 mm [cu (cum unguis: with claw) / su (sine unguis: without claw)], ear 16.5 mm.</p><p>Type locality:  Tiffendell, Eastern Cape Province (30.653°S, 27.928°E), South Africa .</p><p>Paratypes: Ten specimens collected from Tiffendell Ski Resort in December 2013. Some have formaldehydepreserved skins and damaged skulls [TM49094 (BAR1002), male; TM49100 (BAR1016), unknown sex; TM49097 (BAR1019), male; TM49091 (BAR1021), male; TM49095 (BAR1026), male; and TM49098 (BAR1018), unknown sex], one has an intact skull, but the skin was rotten and discarded [TM49092 (BAR1001), female], and three have damaged skulls and rotten skins that were discarded [TM49096 (BAR1006), male; TM49093 (BAR1010), male; and TM49099 (BAR1017), unknown sex]. The specimens were decapitated and the brains removed from some individuals, as part of an earlier study by G. Goldner .</p><p>Referred specimens having molecular identification: (includes the paratypes mentioned above).  From Tiffendell Ski Resort, all collected by G. Goldner in December 2013: TM49099 (BAR1017),  male, unknown age [class 1 (for explanation of relative age/tooth-wear classes, see Taylor et al., 1993)]; TM49098 (BAR1018),  female of unknown age (class 2); TM49092 (BAR1001),  female, unknown age (class 1); TM49094 (BAR1002),  male, unknown age (class 2); TM49096 (BAR1006),</p><p>F -values are indicated for ANOVA tests; all tests were significant at P &lt;0.01 (***). Superscript letters indicate non-significant subsets of means based on pairwise Tukey’s tests. For abbreviations of variables, see under text for Material and Methods.</p><p>male, adult (class 3);  TM49093 (BAR1010), male, adult (class 3);  TM49100 (BAR1016), unknown age (class 1) and sex;  TM49097 (BAR1019), male of unknown age (class 1);  TM49091 (BAR1021), male of unknown age (class 1); and  TM49095 (BAR1026), male of unknown age (class 1). <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=24.936&amp;materialsCitation.latitude=-32.24925" title="Search Plazi for locations around (long 24.936/lat -32.24925)">From Asante Sana</a> (− 32.24925S, 24.936E), 2090 m a.s.l. :   DM13662 (AS 23), skull only of unknown sex, collected 7 June 2009 by <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=25.686&amp;materialsCitation.latitude=-32.521" title="Search Plazi for locations around (long 25.686/lat -32.521)">Armand Kok. From Mountain Zebra National Park</a> (− 32.521S, 25.686E), 1740–1762 ma.s.l  .:   DM13657 (MZ5), skull only, unknown sex, collected 26 July 2009 by  Armand Kok .</p><p>Referred specimens having only morphological identification:   TM22642, 32 km from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=26.7312&amp;materialsCitation.latitude=-31.7696" title="Search Plazi for locations around (long 26.7312/lat -31.7696)">Sterkstroom on Queenstown Road</a>, Eastern Cape Province (− 31.7696S, 26.7312E), adult female, collected by A.J. Prinsloo and J.G. Greeff on 26 July 1957 ;   TM22651, Quqodalo Location, between Glen Grey and <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=26.8823&amp;materialsCitation.latitude=-31.7497" title="Search Plazi for locations around (long 26.8823/lat -31.7497)">Tarkastad</a>, Eastern Cape Province (− 31.7497S, 26.8823E), adult female, collected by J.G. Greeff on 18 October 1955 ;   TM22655, 32 km on road from Sterkstroom to <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=26.7312&amp;materialsCitation.latitude=-31.7696" title="Search Plazi for locations around (long 26.7312/lat -31.7696)">Queenstown</a>, Eastern Cape Province (− 31.7696S, 26.7312E), adult male, collected by A.J. Prinsloo and J.G. Greeff on 26 July 1957 ;   TM22652, Matyhantya No. 11 Location, between Tarkastad and <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=27.0705&amp;materialsCitation.latitude=-31.7825" title="Search Plazi for locations around (long 27.0705/lat -31.7825)">Glen Grey</a>, Eastern Cape Province (− 31.7825S, 27.0705E), adult male, collected by J.G. Greeff on 22 October 1957 ; and   TM29521, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=22.6266&amp;materialsCitation.latitude=-32.2745" title="Search Plazi for locations around (long 22.6266/lat -32.2745)">Karoo National Park</a>, 9 km NW Beaufort West, Western Cape Province (− 32.2745S, 22.6266E), adult female, collected by I.L. Rautenbach, J.G. De Jager and G. De Graaff on 22 January 1979  .</p><p>Incertae sedis: There were two categories of undetermined specimens that were provisionally referred to  O. willani: (1) those specimens without molecular data that were morphologically most like  O. willani (e.g. small-sized in cranial dimensions), albeit slightly distinct (see Fig. 3, marked as ‘  O. cf. karoensis sp. 2’): NMB 9606 and NMB 9607, unknown sex, Glen Agricultural College, Free State Province (− 28.875S, 26.375E); NMB 11420, unknown sex, Sandymount Park, Fauresmith, Free State Province (− 29.625S, 25.125E); and (2), those specimens that most closely resembled  O. willani in morphology but that belonged to a different mtDNA molecular clade (Fig. 2): DM 13661 (MZP 7), female collected at Mountain Zebra National Park (− 32.521S, 25.686E) on 12 February 2010 by Armand Kok.; and DM 13658 (MZ 2), unknown sex, collected at Mountain Zebra National Park (− 32.521S, 25.686E) on 26 July 2009 by Armand Kok.</p><p>Etymology: The species is named for Dr Ken Willan, a South African rodent biologist who conducted pioneering research into the social ecology and reproductive biology of  Otomys and other African rodents. He designed the ‘Willan trap’, which significantly improved the trap success of catching notoriously trap-shy  Otomys (Willan, 1979) .</p><p>Description: Like all members of the genus, it is a relatively large, robust, vole-like rodent, with a large blunt head, short tail and shaggy pelage.  Otomys willani is the smallest member of the genus found in southern Africa, having a mass of 75–83 g (mean 78.2 g, N = 4), total length of 170–198 mm (mean 184 mm, N = 2), tail of 59–74 mm (mean 64.5 mm, N = 4), hindfoot (cu) of 21.5–25.5 mm (mean 23.5 mm, N = 9) and ear 15–20 mm (mean 17.3 mm, N = 4) (data obtained from Phukuntsi et al., 2016: supplementary material). The pelage colour is overall darkish brown with reddish to orange tints, but the facial vibrissae are a pallid creamy-brown colour (Phukuntsi et al., 2016). The upper and lower incisors each have single deep grooves and an additional shallow groove in the lower incisors. The lower M1 has four laminae; upper M3 has seven (in one specimen examined) or six laminae (in ten specimens examined).</p><p>Diagnosis:  Otomys willani can be distinguished in the field from co-occurring  O. auratus and  O. sloggetti within its range in the southern escarpment of South Africa. It is pallid-russet in colour dorsally. Compared with  O. sloggetti, the body is darker reddish in colour, and this species has smaller body measurements than  O. sloggetti . In  O. willani, the pelage is grizzled owing to interspersed black- and russet-coloured hairs (based on the colour of subterminal bands), whereas in  O. sloggetti it is much less grizzled owing to many fewer black hairs intermixed with pale-buff-coloured hairs (Fig. 5). Compared with  O. willani,  O. karoensis has a more pallid-buff rather than pale-russet wash owing to the subterminal section of the guard hairs being buff to pale brown as opposed to russet. Compared with specimens of  O. auratus,  O. irroratus and  O. laminatus, in addition to its distinctly smaller size, the dorsal colour of  O. willani is conspicuously paler that the last-mentioned three species in spite of having a similar hue (Fig. 5).</p><p>Although sample sizes are variable owing to the poor state of preservation of some skins, the series from Tiffendell is instructive, with  O. willani being distinctly smaller than  O. sloggetti in body size without overlap [e.g. mean mass was 78 g (75–83 g, N = 4) in  O. willani and 127 g (116–158 g, N = 5) in  O. sloggetti; total body length was 170–198 mm in two individuals of  O. willani and 224 mm in one  O. sloggetti individual; mean hindfoot (cu) length was 23.5 mm (21.5–25.5 mm, N = 9) in  O. willani and 27.9 mm (26.5–30.5 mm, N = 7) in  O. sloggetti]. The tail of  O. willani is relatively longer (mean 64.5 mm, N = 4, 35% of total length) than in  O. sloggetti (mean 67.5, N = 2, 30% of total length). In the case of co-occurring  O. willani and  O. auratus, as mentioned above, the latter species is much darker buff-brown in colour and also distinctly larger in body and cranial size. In terms of craniodental characters,  O. willani is easily distinguished from  O. sloggetti by having at least one conspicuous groove on the lower incisor (none in  O. sloggetti) and a round-shaped petrotympanic foramen (slit shaped in  O. sloggetti). On craniometric grounds,  O. willani is easily separated from both  O. auratus and  O. irroratus on its smaller size, there being minimal or no overlap in most cranial variables (see Table 3; e.g.  O. willani has GLS &lt;37 mm, MXTL &lt;9 mm and PAL &lt;20 mm, cf. minimal values exceeding the values of these variables in the case of  O. auratus and  O. irroratus). Only in the case of IOC is  O. willani significantly smaller than  O. karoensis (Table 3). As shown by PCA (Fig. 3),  O. willani and  O. karoensis are completely separated on PC3, which is defined as a shape vector. contrasting IOC and PAL (positive loadings) and NAW (negative loadings).  Otomys karoensis generally has a disproportionately large interorbital, longer palatal length (and palatal foramina; Fig. 4) and disproportionately narrow nasal bone relative to  O. willani, reflected in the ratio of IOC/NAW (0.69 in  O. karoensis and 0.66 in  O. willani). In addition to these subtle morphological differences, the two species are phylogenetically distinct from each other based on mtDNA data, and they are not even sister species; instead,  O. karoensis is sister to  O. laminatus (Figs 2, 3). The two species are also geographically and ecologically separated, with  O. karoensis occurring mainly in fynbos biome habitats in the Cape Fold Belt Mountains and  O. willani occurring in grassland biome habitats in the Southern Escarpment.</p><p>Distribution and biology: The species occurs along the Southern Great Escarpment at elevations&gt; 1000 m a.s.l., from the Karoo National Park near Beaufort West in the west to Giant’s Castle in KwaZulu-Natal Province in the east (Fig. 1). Small-sized specimens from two localities on the central plateau of South Africa north of the Drakensberg, referred to as  O. cf. karoensis sp. 2 (dark blue circles in Fig. 1), are provisionally referred here to  O. willani .</p></div>	https://treatment.plazi.org/id/03B887AC26659820FCB148F8FD9EFE75	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Taylor, Peter John;Kearney, Teresa;Dalton, Desire Lee;Chakona, Gamuchirai;Kelly, Christopher M. R.;Barker, Nigel P.	Taylor, Peter John, Kearney, Teresa, Dalton, Desire Lee, Chakona, Gamuchirai, Kelly, Christopher M. R., Barker, Nigel P. (2020): Biomes, geology and past climate drive speciation of laminate-toothed rats on South African mountains (Murinae: Otomys). Zoological Journal of the Linnean Society 189: 1046-1066
03B887AC26789821FF404A13FE13FDAF.text	03B887AC26789821FF404A13FE13FDAF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Otomys karoensis Roberts 1931	<div><p>Otomys cf. karoensis sp. 1 clade</p><p>Vlei rat species show extreme morphological conservatism, which makes them difficult to identify in the field. Small, pallid-coloured specimens from the Sneeuberg Range (preset study) and Tiffendell (from the study by Phukuntsi et al. 2016) were genetically distinct from  O.auratus,  O.irroratus and  O.sloggetti and from  O. karoensis s.s. Morphometric and craniodental analysis (Fig. 3 and Results above) confirm their specific separation from these four species, highlighting the crucial importance of retaining museum voucher specimens in molecular studies, especially in the case of cryptic species. We describe this new  O. cf. karoensis sp. 1 lineage as  O. willani above. This result confirms our hypothesis, based on the cryptic speciation observed in  O. irroratus s.l. (Engelbert et al., 2011), that co-occurring  O. karoensis s.l. would also show evidence of speciation between grassland and fynbos biomes from the Southern Escarpment and Cape Fold Belt mountain ranges, respectively. The drivers of this speciation were therefore probably the same for both species complexes and are described below for the  O. irroratus –  O. auratus clade.</p></div>	https://treatment.plazi.org/id/03B887AC26789821FF404A13FE13FDAF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Taylor, Peter John;Kearney, Teresa;Dalton, Desire Lee;Chakona, Gamuchirai;Kelly, Christopher M. R.;Barker, Nigel P.	Taylor, Peter John, Kearney, Teresa, Dalton, Desire Lee, Chakona, Gamuchirai, Kelly, Christopher M. R., Barker, Nigel P. (2020): Biomes, geology and past climate drive speciation of laminate-toothed rats on South African mountains (Murinae: Otomys). Zoological Journal of the Linnean Society 189: 1046-1066
03B887AC26789822FF404F2EFCFBFCC0.text	03B887AC26789822FF404F2EFCFBFCC0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Otomys irroratus (Brants 1827)	<div><p>Otomys irroratus –  O. auratus clade</p><p>Phylogenetic analysis of Cytb gene sequences reveal the existence of two major evolutionary lineages with no shared haplotypes, consistent with the results of Taylor et al. (2009a) and Engelbrecht et al. (2011). These two lineages, which diverged 1.1 Mya (Taylor et al., 2009a; Engelbrecht et al., 2011), coincide with different biomes and mountain ranges ( O. irroratus with the Cape Fold Belt and the fynbos biome and  O. auratus with the Great Escarpment and the grassland biome). A contact zone between the two lineages was identified at Alice in the Eastern Cape by Engelbrecht et al. (2011). The pattern we observed is mirrored exactly in the case of another montane rodent, the four-striped mouse  Rhabdomys pumilio (Sparrman, 1784) s.l., whose range corresponds closely to that of  O. irroratus s.l., where disjunct genetic lineages coincided with grassland and fynbos biomes and also showed a contact zone in the region of Alice (Du Toit et al., 2012). Likewise, Willows-Munro &amp; Matthee (2011) identified distinct lineages of montane forest shrews from the fynbos and grassland biomes of South Africa. Together with the fact that the  O. karoensis s.l. complex shows exactly the same pattern of vicariance (see above), this striking congruence across several distinct montaneadapted rodent and shrew taxa points to a common evolutionary cause, as detailed below.</p><p>Our new data from AS allow us to extend the western limit of  O. auratus in the Eastern Cape from ~ 27°E (Hogsback) to 25°E (AS). The origin of the two major clades within  O. irroratus s.l. (and  O. karoensis s.l.) might be attributable to vicariance processes caused by aridification. Historically,Africa has experienced periods of high-latitude glaciation cycles, which influenced African climates from the Late Pliocene, causing periods of aridification (Lawes et al., 2007; Taylor et al., 2009a). In southern Africa, this climate change is thought to have caused the onset of drier and/or warmer conditions that led to shrinking of grasslands and fragmentation of biomes owing to increased temperature and decreased rainfall (deMenocal, 2004; Taylor et al., 2009 a, 2015). Aridification of the biomes might have caused the preferred habitat of the species to shrink in size away from other suitable habitats, thereby separating the species distribution into two or more isolated ranges and preventing gene flow between them. A genetic study of snails occurring in the Southern Escarpment demonstrated the importance of aridification in the vicariance of lineages (Barker et al., 2013). Temperateadapted species, such as  O. irroratus and  O auratus, would have been displaced to higher elevations in both the Drakensberg and the Cape Fold Mountain Ranges, effectively isolating populations to these two distinct mountain ranges, whose higher elevations retained temperate climates. Therefore, climatic change linked to elevational heterogeneity is the most likely cause for the distinct difference in the two clades of  O. irroratus s.l. Specifically, the dated split between  O. irroratus and  O. auratus of 1.1 Mya corresponds closely to a period of aridification ~1 Mya (deMenocal, 2004; Taylor et al., 2009a). The fact that the range of  O. irroratus corresponds closely to both the Cape Fold Belt geology and the fynbos biome, whereas the range of  O. auratus corresponds to both the grassland biome and the Great Escarpment mountain system, suggests that although geology combined with climate might have been the cause of the initial divergence of lineages, subsequent speciation and evolutionary divergence might have been aided by ecological coadaptation to the different vegetation biomes.</p><p>From mismatch coefficients from mtDNA sequences, Engelbrecht et al. (2011) showed that  O. irroratus had a unimodal distribution indicative of a recent population expansion, possibly explaining the eastern expansion of  O. irroratus populations away from the Cape Fold Mountains in the Eastern Cape Province to form a secondary contact zone with  O. auratus at Alice on the Great Escarpment. In contrast,  O. auratus has a multimodal mismatch indicative of a stable (and older) population. As also found by Taylor et al. (2009a) and Engelbrecht et al. (2011), our data reveal two distinct subclades in  O. auratus, which are distributed west (central plateau of South Africa and the eastern highlands of Zimbabwe) and east (eastern coastal escarpment populations extending from Alice and Hogsback in the Western Cape to populations from the Drakensberg foothills and midlands of KwaZulu-Natal) of the Great Escarpment. This event was dated at 0.66 Mya by Engelbrecht et al. (2011) and could be explained by a period of warming and/or aridification that resulted in populations becoming fragmented and shrinking downslope on the foothills and escarpments west and east of the high Drakensberg Mountains. Significantly, populations from the Sneeuberg Range are affiliated with the western clade from the Central Plateau rather than with the eastern coastal clade, although populations from Mt Zebra (western clade) and Hogsback (eastern clade) are separated by only 100 km.</p></div>	https://treatment.plazi.org/id/03B887AC26789822FF404F2EFCFBFCC0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Taylor, Peter John;Kearney, Teresa;Dalton, Desire Lee;Chakona, Gamuchirai;Kelly, Christopher M. R.;Barker, Nigel P.	Taylor, Peter John, Kearney, Teresa, Dalton, Desire Lee, Chakona, Gamuchirai, Kelly, Christopher M. R., Barker, Nigel P. (2020): Biomes, geology and past climate drive speciation of laminate-toothed rats on South African mountains (Murinae: Otomys). Zoological Journal of the Linnean Society 189: 1046-1066
03B887AC267B9822FF724DE7FAD7FAD7.text	03B887AC267B9822FF724DE7FAD7FAD7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Otomys sloggetti subsp. clades	<div><p>Otomys sloggetti clades</p><p>Our study reveals new distribution records for  O. sloggetti that extend the range beyond the Drakensberg Range to the adjacent Sneeuberg region of the southern Great Escarpment.  Otomys sloggetti was collected at the three localities that were sampled in the Sneeuberg Range (AS, MZNP and SPNR). However, other mountain regions between the Sneeuberg and Drakensberg, such as the Winterberg and Stormberg, have not been sampled. The existence of three distinct clades of  O. sloggetti from the Sneeuberg, Drakensberg and Maluti Ranges suggests that some vicariance and/or speciation might have occurred between these lineages.</p><p>CONCLUSION</p><p>Our study brings to six the number of  Otomys endemic or near-endemic species occurring in South African mountains (across the Great Escarpment and Cape Fold Belt). The Sneeuberg Centre of Floristic Endemism is clearly also a hotspot of rodent diversity and endemism, with up to three of these endemic  Otomys species co-occurring syntopically. Given the accelerating threats of anthropogenic and climatic change in temperate mountain grassland ecosytems, the conservation status of these montane endemic rodents needs to be reappraised urgently, particularly in the case of the newly described  O. willani, which is restricted to the southern Great Escarpment. Given expected declines in grasslands owing to climate change and the consequent adjustment of  O. auratus from Least Concern to Near Threatened (Taylor et al., 2015; Child et al., 2016), we propose that  O. willani should be classified nationally and globally as Near Threatened under the A4C IUCN criterion.</p></div>	https://treatment.plazi.org/id/03B887AC267B9822FF724DE7FAD7FAD7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Taylor, Peter John;Kearney, Teresa;Dalton, Desire Lee;Chakona, Gamuchirai;Kelly, Christopher M. R.;Barker, Nigel P.	Taylor, Peter John, Kearney, Teresa, Dalton, Desire Lee, Chakona, Gamuchirai, Kelly, Christopher M. R., Barker, Nigel P. (2020): Biomes, geology and past climate drive speciation of laminate-toothed rats on South African mountains (Murinae: Otomys). Zoological Journal of the Linnean Society 189: 1046-1066
