taxonID	type	description	language	source
C46987C9356FB13CBC65F9A398D0C285.taxon	materials_examined	Type locality: Mozambique, Inhambane.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356FB13CBC65F9A398D0C285.taxon	distribution	Distribution: Musser and Carleton (2005) specified the distribution of South African G. cometes as follows: the savanna woodland biome in southern Africa from Pirie Forest (north-west of King William’s Town) in south-eastern Eastern Cape Province of South Africa north through KwaZulu-Natal and Limpopo provinces into eastern Zimbabwe and Mozambique south of the Zambezi River (see references provided by Musser and Carleton 2005). We consider all members of the cometes group (sensu Bryja et al. 2017) as belonging to this species, hence its distribution is wider and also includes populations in the Southern Rift Mountains (including Mt. Gorongosa and Mt. Namuli in Mozambique, Mt. Mulanje in Malawi, and the Chimanimani Mountains on the Zimbabwe – Mozambique border), Eastern Arc Mountains, and coastal forests of Tanzania (including Mafia Island) and southern Kenya. However, we have no genetically confirmed records of this species in South Africa outside Kwazulu-Natal (Fig. 5), and its occurrence in the Eastern Cape province (the Pirie Forest specimens in the collection of AMNH) should be re-analysed genetically.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356FB13CBC65F9A398D0C285.taxon	discussion	Comments: This is, on average, the largest species of the genus (Supporting Information, File S 3). Musser and Carleton (2005) studied the holotype of G. cometes and the other specimens in the type series noted by Thomas and Wroughton (1908); these animals are, on average, larger and have more inflated bullae than those from north of the Zambezi River (but these might represent Grammomys surdaster, as considered here; see below). Ansell (1978) and Ansell and Dowsett (1988) assigned samples from Zambia and Malawi to G. cometes but were also impressed by the chromatic and morphological contrast between them and the holotype from Inhambane. The population from the Pirie Forest was discussed by Taylor et al. (1994) and Taylor (1998), who noted sympatry between G. cometes and G. dolichurus at some localities and at others a gradation of the diagnostic traits usually used to distinguish the two species. Whether those characters have limited discriminatory use in these areas or whether hybridization is occurring is unclear; however, the two species are clearly distinguishable by all genetic markers in our study. We observed that G. cometes has no axial and four inguinal teats (a trait shared with G. dryas, suggesting its ancestral character). This was already used as a discriminating traits by Roberts (1938), who described silindensis (cometes group) and vumbaensis (surdaster group) with differing teat counts: 0 + 4 in the former and 2 + 4 in the latter. The genotyped specimens from the Southern Rift Mountains (especially Malawi) and Eastern Arc Mountains (e. g. Usambara) were reported as G. ibeanus in previous studies (e. g. Stanley and Goodman 2011) and collections (mainly FMNH), but G. ibeanus is not distributed in these mountains (see its account). The short DNA barcode of the type of silindensis belongs to the se 2 lineage, and the holotype and paratype of selousi have mtDNA of se 4 lineage (Supporting Information, File S 1). We therefore propose to synonymize both of them with G. cometes. All available types bearing the names silindensis, selousi, and cometes cluster unequivocally with the genotyped specimens from the cometes group in the geometric morphometric analysis (Table 1). Intriguingly, the type of littoralis was classified as G. cometes, which might represent an extension of its documented distribution in the coastal forests of southern Kenya. The species represents a biogeographical element typical for the coastal forests of eastern Africa (sensu Fayolle et al. 2014). Phylogenomic divergence dating analysis by Mikula et al. (2021) suggested the split of the cometes group (i. e. G. cometes) from the rest of the genus in the Pliocene (3.2 Mya), which agrees with the opinion that most of the coastal forest endemics, including mammals, are palaeoendemics (Burgess et al. 1998). The north – south structuring of the species in mtDNA reflects the fragmented nature of coastal forests, possibly maintained by large rivers flowing to the Indian Ocean (e. g. Rufiji, Zambezi, and Limpopo) or repeated shrinking of coastal forests and their shift to higher elevation owing to climate changes and increases in sea level (Burgess et al. 1998). This suggests that species living in coastal forests colonized parts of the Eastern Arc Mountains and Southern Rift Mountains, probably via riverine gallery forests (see also J. Krásová, O. Mikula, J. Bryja, R. Šumbera, unpublished observations).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356FB13CBC65F9A398D0C285.taxon	biology_ecology	Karyotype (see Fig. 2): 2 n = 49 – 50, FNa (autosomal fundamental number) = 56, FN (fundamental number) = 58 (Denys et al. 2011, as G. selousi). The karyotypes reported as G. dolichurus (2 n = 44; Woodbush Forest Reserve, South Africa; Dippenaar et al. 1983) and G. ibeanus (2 n = 44 – 48; Nyika Plateau, Malawi; Chitaukali et al. 2001) might represent G. cometes. All specimens reported as G. cometes (2 n = 52) by Kryštufek et al. (2008) are G. dolichurus (see Bryja et al. 2017 for more details).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93568B13CBC21FF1B9FCDC13E.taxon	materials_examined	Type locality: South Africa, near Cape Town.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93568B13CBC21FF1B9FCDC13E.taxon	distribution	Distribution: Eastern part of South Africa, from Limpopo Province along coast through KwaZulu-Natal and Eastern Cape provinces to Port Elizabeth (Fig. 5; de Graaff 1981, 1997, Taylor 1998) and Eswatini (A. Monadjem, pers. comm.); the presence in southern Mozambique and, possibly, Zimbabwe should be confirmed. All previous records reported under this name in other parts of Africa belong to other species, mainly to G. surdaster.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93568B13CBC21FF1B9FCDC13E.taxon	discussion	Comments: One syntype skull of dolichurus from the Cape region is very damaged and could not be included in our morphometric analyses; the other syntype is very young, hence it was also not included. The species name has been attributed to many other taxa, and a number of scientific names were synonymized with it (Musser and Carleton 2005). We restrict it to the South African genetic clade, which seems to be different morphologically (Fig. 4; Supporting Information, File S 3), as also suggested by previous studies. For example, Musser and Carleton (2005) mention that ‘ specimens of true dolichurus from South Africa have duller pelage and more inflated bullae than animals from East and West Africa; should these prove to be diagnostic specific differences, the northern populations should be identified as G. surdaster ’. Our genomic analysis supports this view unequivocally. Here, as first proposed by Roberts (1951), we keep only two synonyms of G. dolichurus: baliolus and tongensis. They were both assigned to the dolichurus group with PP =. 2 only, with a higher probability of belonging to the surdaster group (Table 1), but the two groups are overlapping morphologically (Fig. 4), and the genetic material close to both type localities clearly belongs to G. dolichurus (Fig. 6).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93568B13CBC21FF1B9FCDC13E.taxon	biology_ecology	Karyotype: The karyotype of specimens from the Eastern Cape province and from Port St. Johns is 2 n = 52, FN = 62, FNa = 58 (Kryštufek etal. 2008; notethatallspecimensreportedas G. cometes in their karyotypic and CYTB phylogenetic analyses in fact represent G. dolichurus; see Bryja et al. 2017). Dippenaar et al. (1983) reported two different karyotypes from South Africa, 2 n = 52 and 2 n = 44, but the latter is most likely to be G. cometes (see above).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356BB13FBFD9FD8A9FF2C4F3.taxon	materials_examined	Type locality: Malawi, Zomba.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356BB13FBFD9FD8A9FF2C4F3.taxon	distribution	Distribution: Widely distributed from south-western Democratic RepublicoftheCongo (Kikwitarea) andAngolanescarpment, through Zambia to the Albertine Rift Mountains, Eastern Arc Mountains, and Southern Rift Mountains. It is possible that isolated marginal populations occur south of the Zambezi River, such as in Gorongoza National Park in Mozambique or at the border between Mozambique and Zimbabwe (typelocalityof vumbaensis), andincoastalforestsinKenya and Somalia (previously described as caniceps).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356BB13FBFD9FD8A9FF2C4F3.taxon	discussion	Comments: The species has frequently been reported as G. dolichurus in previous studies (Musser and Carleton 2005); however, G. dolichurus is restricted to South Africa (see above). The type of usambarae from northern Tanzania was synonymized with G. macmillani (Musser and Carleton 2005), but no representatives of the macmillani group are distributed in the Usambara Mountains, from where usambarae was described. We genotyped numerous specimens reported by Stanley and Goodman (2011), and they represent either G. cometes (generally larger body size) or G. surdaster (smaller body size). Although we were not able to include the type of usambarae in our morphological analysis, based on the smaller body size and short molar rows, we synonymize usambarae with G. surdaster (see similar opinion by Smithers and Wilson 1979 and Meester et al. 1986, who listed them under G. dolichurus). The populations in more arid coastal forests in Kenya and Somalia have smaller body size and lighter coloration and were described as G. caniceps (Hutterer and Dieterlen 1984). Their mtDNA belongs unequivocally to the surdaster clade (we sequenced several specimens from the type series), as sister to su 11 from the Lake Manyara National Park in northern Tanzania, but unfortunately, we failed to obtain genomic ddRAD data from the type series of G. caniceps because of low DNA quality. The skulls of the holotype and most paratypes of caniceps are classified with the surdaster group, as is the holotype of oblitus from a geographically close Voi locality (Table 1). The type of angolensis was classified into the surdaster group with PP = 0.73, and G. surdaster is the only species documented genetically in Angola (Fig. 6). The type of vumbaensis from eastern Zimbabwe (Vumba and Mount Selinda) was synonymized with G. macmillani (Musser and Carleton 2005), but no representatives of the macmillani group (sensu Bryja et al. 2017) are distributed in this region (see Figs 6, 7). Instead, it might represent the southernmost extension of the range of G. surdaster, because it was classified to this species with PP = 0.84.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356BB13FBFD9FD8A9FF2C4F3.taxon	biology_ecology	Karyotype: Two specimens of G. surdaster from Uluguru Mountains had karyotype 2 n = 42, NFa = 64 (Corti et al. 2005); we sequenced both karyotyped specimens, and they belong to the lineage su 10. This karyotype is likely to differ from that of G. caniceps (2 n = 56, NFa = 64; based on the figure of the karyotype presented by Hutterer and Dieterlen 1984) only by several Robertsonian translocations. The karyotypes of specimens from Somalia (Roche et al. 1984) are variable between and even within individuals, which is caused by polymorphism in Robertsonian translocations and the presence of numerous B chromosomes. However, the number of larger autosomal arms (unlikely to be B chromosomes) is again very similar or even identical to the karyotype of G. surdaster from the Uluguru Mountains (Corti et al. 2005). The karyotypes from Zambia (2 n = 50, NFa = 61 – 62, mtDNA lineage su 7; Corti et al. 2005) and the Katanga region in Democratic Republic of the Congo (2 n = 52, NFa = 62; Matthey 1971, Petter and Tranier 1975) are similar, with slightly lower NFa, which can be explained by a single pericentric inversion.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356BB13EBCC2FA029B0EC57B.taxon	materials_examined	Type locality: ‘ Kenya Colony’ (Uganda), Ruwenzori East, 6000 – 7000 ft (1830 –– 2130 m).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356BB13EBCC2FA029B0EC57B.taxon	distribution	Distribution: A montane species endemic to the Albertine Rift Mountains, i. e. Ruwenzoris and Kivu region in Uganda and Democratic Republic of the Congo, Rwanda, and Burundi (Musser and Carleton 2005). We confirmed it from additional localities in the Albertine Rift, but also from the Minziro Forest (1160 m a. s. l.) in north-western Tanzania and from the Mahale National Park in western Tanzania, where a distinct mtDNA sublineage, m 10, was recorded (Fig. 7).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356BB13EBCC2FA029B0EC57B.taxon	discussion	Comments: Thomas (1907) noted the diagnostic mammary count in G. dryas (no axial pair of teats and only two pairs of inguinal teats), which, in combination with cranial traits, set it apart from other described forms (Musser and Carleton 2005), except G. cometes with the same number of teats (see above). Unfortunately, we were not able to include any representatives of this species in our morphological analysis of skulls. Kerbis Peterhans et al. (1998) reviewed its elevational distribution in the context of the entire Ruwenzori small mammal fauna. We recently collected it close to the type locality in the Rwenzori Mountains (Nyakelengija gate, 1720 m a. s. l.; Supporting Information, File S 1). It is a well-characterized taxon genetically for both mtDNA (having three sublineages with parapatric distribution; Fig. 7 C) and ddRAD data (wherein a single MOTU was identified by both branch-cutting and INFOMAP approaches; Figs 3, 7).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356BB13EBCC2FA029B0EC57B.taxon	biology_ecology	Karyotype: Not known.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356AB121BFC4FDFE9BCBC490.taxon	materials_examined	Type locality: Ethiopia, north of Lake Rudolf (= Turkana), Wouida, elevation 6200 feet (1890 m a. s. l.).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356AB121BFC4FDFE9BCBC490.taxon	distribution	Distribution: The belt of a mosaic of Guineo-Congolian forests and Sudanian savannah, from Sierra Leone to south-western Ethiopia and western Kenya (Fig. 7). In the central parts of the Sudanian savannah, it has been reported as G. dolichurus or G. macmillani (e. g. GBIF database; https: // www. gbif. org /), but it is not confirmed genetically. In Uganda, it is the most widespread species, recorded even in Bugala Island in Lake Victoria (Fig. 7).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356AB121BFC4FDFE9BCBC490.taxon	discussion	Comments: This species corresponds to INFOMAP MOTU VI, encompassing West African mtDNA clade m 3 (formerly called buntingi) and Central and East African mtDNA clade m 5 (Fig. 3). The two clades are sister in both mtDNA and ddRAD phylogenetic analysis; in our view, they simply represent intraspecific (phylogeographical) genetic variation. A similar pattern of west – east differentiation has been found in numerous rodent taxa living at the northern margin of Guineo-Congolian forests and in Sudanian savannah (e. g. Mus musculoides, Temminck, 1853, Bryja et al. 2014; Mastomys erythroleucus (Temminck, 1853), Brouat et al. 2009; Arvicanthis niloticus (Desmarest, 1822), Bryja et al. 2019 a; Lemniscomys zebra (Heuglin, 1864), Hánová et al. 2021; and Gerbilliscus giffardi (Wroughton, 1906), Granjon et al. 2012). There are many names previously used for Grammomys in this geographical range (Fig. 7). The oldest name is macmillani, described from Wouida, north of Lake Rudolf (= Turkana). This area can be inhabited also by the mtDNA clade m 4 (INFOMAP MOTU IV, found in well-grown forests in higher elevations, from the Kenyan highlands to south-western Ethiopia, considered here as G. ibeanus; see below). However, the macmillani type instead represents the widespread species in the belt of Sudanian savannah. Firstly, the type locality is not described precisely. Based on the species description (Wroughton 1907), the holotype was collected by the expedition of Mr Ph. C. Zaphiro on 30 June 1905 in Wouida, north of Lake Rudolf, elevation 6200 ft (1890 m a. s. l.). We were unable to identify habitats in ‘ Wouida’, and we did not capture any specimens with m 5 mtDNA in southern Ethiopia, but the specimen from this mitochondrial clade from the same elevation in South Sudan clustered unequivocally with MOTU VI in all ddRAD analyses. Many other taxa typical of Sudanian savannah were recently documented from drier woodlands in south-western Ethiopia (e. g. Gerbilliscus giffardi or Aethomys hindei (Thomas, 1902); Bryja et al. 2019 b) and north-western Uganda (S. W. Babyesiza, J. Bryja, pers. observation). The presence of this Grammomys clade is therefore very likely in drier woodlands in the region ‘ north of Lake Rudolf ’, i. e. the type locality of G. macmillani, which also represents its easternmost record. Second, two specimens from Chak Chak (now South Sudan) are mentioned to be almost identical with the type of macmillani from Wouida in the description by Wroughton (1907). They were later described as G. macmillani gazellae by Thomas (1910), but the investigation by Hutterer and Dieterlen (1984) again confirmed that the types of macmillani and gazellae are virtually identical. Spermatozoal morphology was documented by Breed (1995; as gazellae). The reports of G. macmillani in the Eastern Arc Mountains of Tanzania, documented by Stanley et al. (1998), represent another taxon (most likely MOTU VII, for which we propose the name G. polionops; see below). Setzer (1956) suggested (without any genetic data) that none of the characters formerly used to separate G. macmillani and G. surdaster is of more than subspecific value, and he proposed that the animals formerly known as surdaster should henceforth be called macmillani. Our morphological analysis suggests high similarity of these taxa (Fig. 4; Table 1), but they are distinct both genetically and geographically. Based on the locations of type localities and distributions of the genomically well-delimited surdaster and macmillani groups (sensu Bryja et al. 2017), we agree with Musser and Carleton (2005) and place in synonymy with G. macmillani the following names: callithrix, gazellae, and erythropygus. Furthermore, discolor, elgonis, and insignis were synonymized with G. dolichurus (= surdaster in the work of Setzer 1956) but should also be here listed as synonyms of G. macmillani, based on the locations of their type localities. The taxon aridulus was listed either as a subspecies of G. macmillani (Allen 1939, Ellerman 1941, Setzer 1956) or included in G. dolichurus (Misonne 1974) but was considered a distinct species by Hutterer and Dieterlen (1984). Based on its description, it represents ‘ an unusually pallid, desert-coloured species … skull resembles G. buntingi from Liberia … In outward appearance it is perhaps more like G. s. elgonis than any other, while in skull it resembles the outwardly very different G. buntingi. ’. We obtained a short CYTB sequence from two specimens of the type series collected on Mt. Jebel Marra in Sudan, and they clearly cluster with m 5. We therefore propose to synonymize aridulus with G. macmillani, which also makes sense from a biogeographical perspective (Fig. 7). Except for Misonne (1974), who included it in G. dolichurus, buntingi has always been listed or discussed as a distinct species in all recent reviews (Petter and Trainer 1975, Hutterer and Dieterlen 1984, Musser and Carleton 2005, Kryštufek 2008, Denys et al. 2011). The western African population forms a separate subclade in both mtDNA (Fig. 2) and ddRAD (Fig. 3) phylogenies, but INFOMAP clustered them together with central and eastern African populations into MOTU VI (Fig. 3). Morphologically, these populations are very similar, differing by very small details of the teeth (Petter and Tranier 1975), and we were not able to find any clearly distinguishing traits compared with populations of Grammomys from western Africa and the Central African Republic in the MNHN collection. Based on the genetic and morphological similarity, we therefore propose to synonymize buntingi with G. macmillani and consider it as an intraspecific phylogeographical lineage.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C9356AB121BFC4FDFE9BCBC490.taxon	biology_ecology	Karyotype: Karyotype was analysed by Petter and Tranier (1975; as buntingi and gazellae) and Civitelli et al. (1989; as gazellae). The population of buntingi from Cote d’Ivoire has 2 n = 52, NFa = 58 (Peter and Tranier 1975). The karyotype of gazellae from Central African Republic is highly variable because of the presence of numerous B chromosomes, but if they are excluded, then 2 n = 54, NFa = 58 (based on fig. 2 of Civitelli et al. 1989), i. e. they differ by only a single Robertsonian translocation, supporting their conspecificity.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93575B121BF8DFC599E76C4B1.taxon	materials_examined	Type locality: Kenya, Molo.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93575B121BF8DFC599E76C4B1.taxon	distribution	Distribution: Montane forests in Kenyan Highlands, eastern Uganda, South Sudan, south-western Ethiopia, and Mt. Kilimanjaro.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93575B121BF8DFC599E76C4B1.taxon	discussion	Comments: Hutterer and Dieterlen (1984) treated ibeanus as a form of G. cometes (both taxa are relatively large-bodied), but the striking morphological distinctions between samples of ibeanus and the type series of cometes analysed by Musser and Carleton (2005) prompted specific ranking of ibeanus. We include in G. ibeanus two INFOMAP gene pools: MOTU IV from montane forests in Kenyan Highlands, eastern Uganda (Mt. Elgon and Mt. Morungole), the Imatong Mountains in South Sudan, highlands in the Ilemi Triangle, and forests of south-western Ethiopia, and MOTU V from Mt. Kilimanjaro. Both MOTUs are sister clades in both mitochondrial and genomic trees (albeit in the former with only weak support; Fig. 2). The species can be sympatric with G. macmillani (Imatong Mountains, north-eastern Uganda, and western Kenya) and G. polionops (Nanyuki, Kenya), but the sympatric populations are genetically well distinguishable in both mtDNA and nuDNA (Fig. 7), supporting their reproductive isolation. The ecological differences, e. g. the preference for different types of habitats and elevation, are worth further studies. The population from Mt. Kilimanjaro (MOTU V) is highly distinct genetically (Fig. 3) and separated geographically from all other populations of G. ibeanus. More detailed morphological and ecological studies in the future might possibly lead to its full species status (and proposal of a new name, given that no Grammomys was described from Mt. Kilimanjaro). The specimens in FMNH identified as G. ibeanus from the Eastern Arc Mountains in Tanzania, from Malawi, and from Mozambique belong genetically to the cometes group (sensu Bryja et al. 2017) and represent G. cometes (as defined in this study; see above). The taxon lutosus (from Mt. Nyiro in Kenya) was listed as a subspecies of G. ibeanus by Allen (1939). In our morphological analysis, it was classified into the surdaster group (PP = 0.67), but the holotype is a sub-adult specimen with a broken skull, and the analysis was based on only a subset of landmarks. Geographically, it might belong to G. ibeanus, hence we keep it as its younger synonym. The holotype of T. gigas from Mt. Kenya in our opinion simply represents an extremely large individual of G. ibeanus (see also Musser and Carleton 2005, Denys et al. 2011). All recently genotyped specimens from Mt. Kenya (collected by K. Onditi and colleagues) have mtDNA of the lineage m 4 (Supporting Information, File S 1), supporting this conclusion. The holotype of gigas was classified as a member of the cometes group (with PP = 0.95; Table 1), which is attributable to its large size. When only shape is considered, it fits in the macmillani group (PP = 0.90).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93575B121BF8DFC599E76C4B1.taxon	biology_ecology	Karyotype: Not known.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93575B120BCDFFC389BDCC4B1.taxon	materials_examined	Type locality: Lukenya Mt., Ulukenia Hills, Kenya.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93575B120BCDFFC389BDCC4B1.taxon	distribution	Distribution: Eastern Arc Mountains (e. g. Udzungwa and a few other hills; not in Usambara), the Gregory Rift in northern Tanzania and southern Kenya, and the southern part of the Great Rift Valley in Ethiopia.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93575B120BCDFFC389BDCC4B1.taxon	discussion	Comments: The name polionops is the oldest available name for populations grouped in the INFOMAP MOTU VII and branch-cutting MOTU 6 (Figs 3, 7 D). The species inhabits relatively dry forests and woodlands along the Gregory branch of the East African Rift. At one locality (north of Nanyuki, Kenya), the two mtDNA lineages of G. polionops (m 7 and m 8) were found together with G. ibeanus (mtDNA lineage m 4), but the two species were clearly distinguishable by ddRAD data, suggesting a reproductive barrier between these two species. In the Eastern Arc Mountains and Northern Highlands of Tanzania and southern Kenya, the species can be sympatric with G. surdaster, and the mechanisms of their co-occurrence require further studies (e. g. how they differ morphologically and ecologically and whether they can hybridize). The taxon from the vicinity of Arba Minch in Ethiopia was described as G. minnae, based on karyotype (Hutterer and Dieterlen 1984), but genomic data were not able to distinguish it from the populations in central and southern Kenya. Both the holotype and the paratype of minnae were clearly classified to the macmillani group by the morphometric analysis. The holotype of brevirostris, described from Loita Plains by Kryštufek (2008), was sequenced at CYTB, and it belongs to the mitochondrial lineage m 8, clustering with the genotyped specimens from the nearby Loita Hills (Supporting Information, File S 1). The morphologically distinct holotype of brevirostris might represent a typical individual of G. polionops, because no individuals assigned here to G. polionops were included in Kryštufek’s (2008) morphological comparison. The holotype of brevirostris is assigned to the macmillani group (that also includes G. polionops) with PP = 0.85 (Table 1).	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
C46987C93575B120BCDFFC389BDCC4B1.taxon	biology_ecology	Karyotype: Three very different karyotypes (but all of them with low diploid numbers; Fig. 2) were described for populations that we assign to G. polionops. The role of such different karyotypes in reproductive isolation should be explored further. The specimen from northern Tanzania (a male from Jipe, no. T 50485, belonging to mtDNA lineage m 7) had 2 n = 20, NFa = 31 (Corti et al. 2005). Another specimen from northern Tanzania (a female from Ndaleta, ind. no. TZ 12, with mtDNA of m 8) had 2 n = 27 and NFa = 39 (Fadda et al. 2001). Finally, the taxon minnae from the Bulcha Forest in Ethiopia was described based on the karyotype 2 n = 32 and NF = 64 (figured by Olert et al. 1978). The same karyotype was found recently in specimens from the Arero forest in southern Ethiopia (L. Lavrenchenko, unpub. obs.) with mtDNA of the lineage m 11.	en	Bryja, Josef, Kerbis Peterhans, Julian C., Lavrenchenko, Leonid A., Nicolas, Violaine, Denys, Christiane, Bryjová, Anna, Šumbera, Radim, Mikula, Ondřej (2025): Integrative taxonomic revision of the African thicket rats (Murinae: Grammomys): how genomics decreases the number of currently recognized species. Zoological Journal of the Linnean Society (Statistics in Society) 203 (2): 1-20, DOI: 10.1093/zoolinnean/zlae057, URL: https://doi.org/10.1093/zoolinnean/zlae057
