Pseudoromicia kityoi, 2020

Monadjem, Ara, Demos, Terrence C, Dalton, Desire L, Webala, Paul W, Musila, Simon, Kerbis Peterhans, Julian C & Patterson, Bruce D, 2020, A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species, Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 190, pp. 1-33 : 20-23

PSEUDOROMICIA KITYOI MONADJEM, KERBIS PETERHANS, NALIKKA, WASWA, DEMOS & PATTERSON SP. NOV.

KITYO’ S SEROTINE

LSID: http://zoobank.org/ urn:lsid:zoobank.org:pub:71737F08-2938-4403-8385-5438B2E5EABE

Holotype: FMNH 223211 , field number JCK 7436 . This specimen was collected by Betty Nalikka and Sadic Waswa Babyesiza during a field training exercise with Julian Kerbis Peterhans. It is an adult male preserved in ethanol, with skull extracted and cleaned, and tissue taken from breast muscle and preserved in dimethyl sulfoxide. Type locality: Mabira Forest Reserve, 0.79 km northeast of Nagojje Station , Mukono District of the Central Region , Uganda; geographical coordinates: 0.4451°N, 32.88876°E ( Fig. 1 ). The type specimen was netted on 19 October 2012 in cultivated gardens directly adjacent (for a photograph of the type locality, see Fig. S2 ) to Mabira forest at an elevation of 1130 m above sea level .

Paratype: One other male ( FMNH 223555 ) was netted at the same location and on the same night as the holotype, and closely resembles it genetically ( Fig. 3B ) and morphologically ( Tables 5–7) and can therefore be considered a paratype.

Etymology: This species is named in honour of Dr. Robert M. Kityo, mammalogist, mentor and longserving curator at the Museum of Zoology, Makerere University, in recognition of his valuable contributions to bats and small mammal research in the region. His welcoming nature, curiosity, hospitality and support have facilitated numerous and diverse research agendas over the decades for both national and international researchers.

Diagnosis: This is the largest member of the genus Pseudoromicia , with forearm length of 37 and 38 mm ( Table 5) and greatest skull length of 14.70 and 14.99 mm for the two known specimens ( Table 6). In comparison, the maximum greatest skull length in Pse. roseveari (which is the second largest member of the genus) is 14.5 mm ( Table 6). Pseudoromicia brunnea is smaller in forearm length and in most craniodental measurements. Therefore, this species is readily diagnosable by size alone. It can easily be distinguished from the white-winged members of this genus ( Pse. rendalli , Pse. isabella and Pse. tenuipinnis ) by its dark wings.

Description: External characters: Pseudoromicia kityoi is a large-sized pipistrelle-like bat, similar in size to the largest members of the Nycticeinops group, specifically Nyc. macrocephalus and Nyc. happoldorum , which were both described in the genus Paraphypsugo ( Hutterer & Kerbis Peterhans, 2019; Hutterer et al., 2019). Despite its large size, this species is similar in external features to other black-winged members of Pseudoromicia . The pelage is medium brown above and slightly paler below. The individual hairs are unicoloured on the upper parts and bicoloured on the under parts, with the proximal half darker than the distal half. Like Pse. brunnea and Pse. roseveari , the patagium and uropatagium are both dark in colour. The ears are short and rounded, and the tragus has a curved outer margin as is typical of the genus ( Monadjem et al., 2013).

Craniodental characters: The skull is robust for a Pseudoromicia , even more so than in Pse. roseveari . The rostrum has a shallow depression, and the brain case is moderately inflated as in other members of the genus. There is no occipital ‘helmet’ as seen in the cranium of Lae. capensis ( Monadjem et al., 2020b) . The sagittal and lambdoidal crests are visible, and the zygomatic arches are robust for a pipistrelle-like bat ( Fig. 9 ). The dentition in Pse. kityoi is typical of the genus, with I 2/3, C 1/1, P 1/3, M2/3. In the upper tooth row, I 1 is unicuspid and I 2 is tiny, extending slightly beyond the cingulum of I 1. The P 1 is absent, putting C in contact with P 2. The m 3 is myotodont sensu Van Cakenberghe & Happold (2013).

Biology: Owing to the paucity of specimens, almost nothing can be said about the biology of this species. The only two known specimens were captured within 200 m from the edge of Mabira Forest in a domestic garden (Supporting Information, Fig. S2 ). However, considering that most members of this genus are restricted to tropical rainforest habitats, and that the two known specimens of this species were captured in a remnant patch of rainforest, its global distribution might be both fragmented and limited in extent. Urgent surveys are required to assess the status of this species at Mabira Forest Reserve, which has been steadily losing habitat to agriculture over the past few decades ( Boffa et al., 2008). We suggest that this species might be present in other Congo Basin forest patches in Uganda (e.g. Semliki, Kibale, Kashyoha-Kitomi) and Kenya (Kakamega), although extensive surveys at Kakamega forest have failed to locate this species there ( Webala et al., 2019). Owing to the limited information available on this species, we recommend that it be given the IUCN conservation status of ‘Data Deficient’, but we note that because of its presumed close association with rapidly disappearing forest habitat, this species is probably of conservation concern.

Its closest known relative is Pse. roseveari , recently described from Mount Nimba and with a limited distribution in the borderland zone between Liberia and Guinea ( Monadjem et al., 2013; Decher et al., 2015; Mamba et al., in press), some 4700 km to the west. Whether either species occurs in the vast tropical rainforests between these two sites is unknown and deserves investigation.

Figures

Figure 1. Type localities of taxa of African and Malagasy Vespertilionini and Pipistrellini. Valid species are denoted by filled circles, subspecies and synonyms by open circles and species described herein by stars: 1, Pipistrellus abaensis J. A. Allen, 1917; 2, N[ycticejus]. adovanus Heuglin, 1877; 3, Pipistrellus aero Heller, 1912; 4, Vespertilio pipistrellus var. africanus Rüppell, 1842; 5, Nycticeius africanus G. M. Allen, 1911; 6, Scotoecus albigula Thomas, 1909; 7, Scoteinus schlieffeni albiventer Thomas & Wroughton, 1908; 8, Scotophilus albofuscus Thomas, 1890; 9, Vesperugo anchietae Seabra, 1900; 10, Laephotis angolensis Monard, 1935; 11, Eptesicus capensis angolensis Hill, 1937; 12, Pipistrellus ariel Thomas, 1904; 13, Scotoecus artinii De Beaux, 1923; 14, Eptesicus ater J. A. Allen, 1917; 15, Pipistrellus nanus australis Roberts, 1913; 16, Scoteinus schlieffeni australis Thomas & Wroughton, 1908; 17, Scoteinus schlieffeni bedouin Thomas & Wroughton, 1908; 18, Pipistrellus eisentrauti bellieri De Vree, 1972; 19, Hypsugo bemainty Goodman et al., 2015; 20, Vesperus bicolor Bocage, 1889; 21, Laephotis botswanae Setzer, 1971; 22, Pipistrellus (Romicia) kuhlii broomi Roberts, 1948; 23, Vesperugo (Vesperus) brunneus Thomas, 1880; 24, Vespertilio capensis A. Smith, 1829; 25, Scotoecus cinnamomeus Wettstein, 1916; 26, Pipistrellus crassulus Thomas, 1904; 27, Pipistrellus culex Thomas, 1911; 28, Vesperus damarensis Noack, 1889; 29, Scotophilus darwini Tomes, 1859; 30, Pipistrellus deserti Thomas, 1902; 31, Pipistrellus eisentrauti Hill, 1968; 32, Scotoecus falabae Thomas, 1915; 33, Eptesicus faradjius J. A. Allen, 1917; 34, Scoteinus schlieffeni fitzsimonsi Roberts, 1932; 35, Pipistrellus fouriei Thomas, 1926; 36, Pipistrellus kuhlii fuscatus Thomas, 1901; 37, Pipistrellus fuscipes Thomas, 1913; 38, Eptesicus garambae J. A. Allen, 1917; 39, Vespertilio capensis gracilior Thomas & Schwann, 1905; 40, Vesperugo (Vesperus) grandidieri Dobson, 1876; 41, Vesperus guineensis Bocage, 1889; 42, Pipistrellus hanaki Hulva & Benda, 2004; 43, Parahypsugo happoldorum Hutterer, Decher, Monadjem & Astrin, 2019; 44, Pipistrellus helios Heller, 1912; 45, Vespertilio hesperida Temminck, 1840; 46, Scotoecus hindei Thomas, 1901; 47, Scotophilus hirundo de Winton, 1899; 48, Vesperus humbloti Milne-Edwards, 1881; 49, Vesperugo hypoleucus Heuglin [in Fitzinger & Heuglin], 1866; 50, Pipistrellus inexspectatus Aellen, 1959; 51, Neoromicia isabella Decher, Hutterer & Monadjem, 2016; 52, Laephotis kirinyaga Monadjem et al., this paper; 53, Pseudoromicia kityoi Monadjem et al., this paper; 54, Hypsugo lanzai Benda, Al-Jumaily, Reiter & Nasher, 2011; 55, Pipistrellus leucomelas Monard, 1932; 56, Parahypsugo macrocephalus Hutterer & Kerbis Peterhans, 2019; 57, Vesperugo maderensis Dobson, 1878; 58, Eptesicus somalicus malagasyensis Peterson, Eger & Mitchell, 1995; 59, Vespertilio marginatus Cretzschmar, 1830; 60, Pipistrellus marrensis Thomas & Hinton, 1923; 61, Vespertilio matroka Thomas & Schwann, 1905; 62, Pipistrellus africanus meesteriKock,2001;63,Eptesicus melckorumRoberts,1919;64,Scotophilusminimus Noack,1887;65,Pipistrellus minusculus Miller, 1900; 66, Vespertilio minuta Temminck, 1840; 67, Pipistrella minuta Loche, 1867; 68, Pipistrellus musciculus Thomas, 1913; 69, Laephotis namibensis Setzer, 1971; 70, Pipistrellus nanulus Thomas, 1904; 71, Vespertilio nanus Peters, 1852; 72, Eptesicus capensis nkatiensis Roberts, 1932; 73, Scabrifer notius G. M. Allen, 1908; 74, Pseudoromicia nyanza Monadjem et al., this paper; 75, †Scotoecus olduvensis Gunnell, Butler, Greenwood & Simmons, 2015; 76, Vesperugo pagenstecheri Noack, 1889; 77, Pipistrellus (Pipistrellus) permixtus Aellen, 1957; 78, Eptesicus phasma G. M. Allen, 1911; 79, Vespertilio pipistrellus Schreber, 1774; 80, Vespertilio platycephalus Temminck, 1832; 81, Vesperugo pulcher Dobson, 1875; 82, Vesperugo pusillulus Peters, 1870; 83, Pipistrellus raceyi Bates et al., 2006; 84, Eptesicus rectitragus Wettstein, 1916; 85, Vesperugo (Vesperus) rendalli Thomas, 1889; 86, Neoromicia robertsi Goodman et al., 2012; 87, Neoromicia roseveari Monadjem et al., 2013; 88, V[espertilio]. rueppelii J. Fischer, 1829; 89, Scotophilus rusticus Tomes, 1861; 90, Vespertilio savii Bonaparte, 1837; 91, Nycticejus schlieffenii Peters, 1859; 92, Pipistrellus rueppelli senegalensis Dorst, 1960; 93, †Nycticeinops serengetiensis Gunnell et al., 2015; 94, Pipistrellus simandouensis Monadjem et al., 2020; 95, Vespertilio minutus somalicus Thomas, 1901; 96, Vesperugo stampflii Jentink, 1888; 97, Neoromicia stanleyi Goodman et al., 2017; 98, Vesperus tenuipinnis Peters, 1872; 99, Eptesicus ugandae Hollister, 1916; 100, Neoromicia vansoni Roberts, 1932; 101, Pipistrellus vernayi Roberts, 1932; 102, Laephotis wintoni Thomas, 1901; 103, Scotoecus woodi Thomas, 1917; 104, Eptesicus zuluensis Roberts, 1924. Not mapped: [Pipistrellus Kuhli] latastei Laurent, 1937; Vespertilio pusillus LeConte, 1857; Vesperugo subtilis Sundevall, 1846.

Figure 2. Maximum likelihood phylogeny of intergeneric relationships of mitochondrial cytochrome b sequences of Vespertilionidae.The phylogeny was inferred in IQ-TREE, and its topology was similar to the Bayesian phylogeny calculated in MRBAYES. Bootstrap (BS) values followed by Bayesian posterior probabilities (PP) are indicated adjacent to nodes (those nodes with both BS <70% and PP <0.95 are not labelled).

Figure 3. Maximum likelihood phylogeny of mitochondrial cytochrome b sequences of Vespertilionidae: (A) Pipistrellus, Scotoecus, Vansonia, and outgroups (B) Afronycteris, Pseudoromicia, Nycticeinops, and Hypsugo (C) Laephotis and Neoromicia. The phylogeny was inferred in IQ-TREE, and its topology was similar to the Bayesian phylogeny calculated in MRBAYES. Filled red circles on nodes denote bootstrap (BS) values ≥ 70% and Bayesian posterior probabilities (PP) ≥ 0.95. Open circles outlined in black indicate BS ≥ 70% and PP <0.95, and open circles outlined in red indicate BS <70% and PP> 0.95. Support values for most minor clades are not shown. Specimen localities include counties for Kenya. DRC refers to Democratic Republic of the Congo and CAR to Central African Republic. Museum acronyms are defined in the Material and Methods section. Sequences downloaded from GenBank are indicated by inclusion of GenBank accession numbers (Supporting Information, Table S1). Branch colours indicate individual species/clade membership.

Figure 9. Plate showing the cranium and mandible of Pseudoromicia kityoi (FMNH 223211).Scale bar = 5 mm.

Tables

Table 5. External measurements (in millimetres) and mass (in grams) of Pseudoromicia kityoi from Mabira Forest Reserve, Uganda and Pseudoromicia nyanza from Kisumu, Kenya

Specimen or taxon	Total length	Tail length	Hindfoot length	Ear length	Forearm length	Body mass
Pseudoromicia kityoi	88	34	10	10	38.0	7.9
Holotype FMNH 223211
Pseudoromicia kityoi	89	36	9	10	37.0	8.0
Paratype FMNH 223555
Pseudoromicia roseveari	87.3 ± 2.95, 83–93,	39.7 ± 2.65, 34–44,	9.9 ± 1.10, 8–11,	12.9 ± 0.64, 12–14,	36.5 ± 1.54, 32.6–38.0,	6.5 ± 0.50, 6.0–7.0,
	N = 10	N = 9	N = 10	N = 8	N = 10	N = 10
Pseudoromicia brunnea	84.8 ± 3.14, 80–89,	35.9 ± 1.55, 33–38,	7.79 ± 0.97, 7–10,	12.4 ± 0.83, 11–14,	34.6 ± 1.06, 32.8–36.7,	6.0 ± 1.04, 4.8–9.4,
	N = 15	N = 15	N = 14	N = 15	N = 15	N = 15
Pseudoromicia nyanza	83	30	7	11	31.0	8.3
Holotype FMNH 215626
Pseudoromicia nyanza	83.5 ± 2.73, 79–89,	33.3 ± 2.29, 30–38,	7.5 ± 0.66, 7–9,	13.0 ± 0.64, 11–13,	31.2 ± 1.24, 29.0– 33.0,	6.1 ± 0.99, 4.8–8.3,
(other specimens)	N = 13	N = 13	N = 13	N = 13	N = 13	N = 13
Pseudoromicia rendalli	88.5 ± 3.92, 82–95,	37.0 ± 3.15, 32–42,	7.5 ± 0.90, 6–9,	11.3 ± 1.57, 9–13,	34.8 ± 1.30, 33.4–37.0,	5.8 ± 0.94, 4.0–6.8,
	N = 15	N = 9	N = 8	N = 9	N = 9	N = 6
Pseudoromicia isabella	78.8 ± 2.73, 74–82,	29.3 ± 4.23, 24–36,	6.9 ± 1.18, 5–8,	12.3 ± 1.37, 10–14,	30.6 ± 1.47, 28.0– 31.9,	4.7 ± 0.87, 4.0–6.0,
	N = 15	N = 6	N = 6	N = 6	N = 6	N = 6
Pseudoromicia tenuipinnis s.s.	74.2 ± 2.17, 72–77,	29.2 ± 0.84, 28–30,	6.5 ± 0.80, 5–7,	12.7 ± 0.91, 12–14,	29.4 ± 1.18, 28.1– 32.0,	4.0 ± 0.29, 3.5–4.4,
(West Africa)	N = 5	N = 5	N = 7	N = 7	N = 6	N = 6

Measurements are presented as the mean ± SD, range and sample size (N). Measurements are of the holotypes, other individuals of the two new species and other species of Pseudoromicia . The three species listed above the horizontal black line are dark winged, the four below are white winged (see main text for more details).