Laephotis kirinyaga, 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 : 15-18

LAEPHOTIS KIRINYAGA MONADJEM, PATTERSON, WEBALA & DEMOS SP. NOV.

EAST AFRICAN SEROTINE

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

Synonymy

Eptesicus capensis Kingdon (1974) .

Pipistrellus capensis garambe Thorn & Kerbis Peterhans (2009) (in part).

Neoromicia capensis Patterson & Webala (2012) .

Neoromicia somalica Benda et al. (2016) (in part).

Holotype: FMNH 234558 , field number BDP 7516. This specimen was collected by Bruce D. Patterson, Paul W. Webala, Carl W. Dick and Beryl Makori. It is an adult male, with muscle tissue in liquid nitrogen, the body fixed in formalin and preserved in ethanol, now with skull extracted and cleaned. Type locality: Marsabit National Park, 1.3 km SE of campground near Headquarters , Marsabit County, Kenya (2.3090°N, 38.0001°E; Fig. 1 ). The type specimen was netted on 27 July 2015 at an elevation of 1280 m above sea level .

Paratypes: One other male ( FMNH 234559 ) was captured at the same location and on the same night as the type specimen and is considered a paratype. Seven other individuals ( FMNH 234546 , FMNH 234549– 234553 , FMNH 234556–234557 , four males and three females), were collected close to the type locality at elevations ranging from 1157 to 1356 m from 16 to 26 July 2015 (Supporting Information, Table S1 ); they closely resemble the holotype genetically ( Fig. 3C ) and morphologically ( Tables 2–4 ) and are also considered paratypes.

Etymology: The specific epithet is a Kikuyu word for Mount Kenya and reflects the distribution of the species in the northern highlands of Kenya. It is a noun in apposition.

Diagnosis: This species is similar in size and appearance to its sister species Lae. capensis . It is easily distinguished from the long-eared Laephotis species by its shorter ears. Of the short-eared Laephotis species, Lae. stanleyi and Lae. robertsi are significantly larger in forearm length and most craniodental measurements ( Tables 2–4 ). In contrast, Lae. malagasyensis is smaller, especially in cranial measurements ( Table 3 ). Laephotis matroka is similar in external and craniodental measurements but is typically darker brown above and medium brown below ( Goodman, 2011). In any case, Lae. robertsi , Lae. malagasyensis and Lae. matroka are all endemic to Madagascar ( Goodman et al., 2012, 2017) and genetically distinct from Lae. kirinyaga ( Fig. 3C ). Laephotis kirinyaga closely resembles Lae. capensis , from which it differs by 8.3% on the Cytb gene (Supporting Information, Table S4). Externally, the two species are alike and broadly overlap in size, but Lae. kirinyaga is on average smaller in most measurements, particularly total length and forearm length ( Table 2 ). Likewise, Lae. capensis is on average larger for all craniodental measurements (but with significant overlap), except greatest skull height, which is greater in Lae. kirinyaga . This is borne out in the lateral profile of the skull ( Fig. 7 ), which is visibly flatter in Lae. capensis . These two species occupy mostly separate regions in multivariate space ( Fig. 6 ), but again with some overlap. In contrast, the three specimens assigned to Lae. kirinyaga from Ethiopia and Guinea (labelled ‘cf. capensisW’ in Fig. 6 ), for which genetic data are lacking, fall completely within the multivariate space of the Lae. kirinyaga specimens (labelled ‘cf. capensis’) that have been sequenced.

Description: Exernal characters: Laephotis kirinyaga is a medium-sized pipistrelle-like bat, with strongly contrasting fur dorsally and ventrally. The dorsal pelage is medium brown, with most individual hairs being tipped light yellowish brown, giving the bat a brightly coloured appearance. The ventral pelage is cream–white to light cream–brown, with a dark base. The ears are short and rounded, and the tragus is curved distally on both anterior and posterior margins,

ending in a rounded tip, as in Lae. capensis ( Monadjem et al., 2020b) . The ears and muzzle are dark brown in colour, and the skin around the eyes is dark brown in the type specimen ( Fig. 8A ) but mostly pinkish in the paratypes.

Craniodental characters: The skull is relatively robust, as in Lae. capensis , but less so than in Lae. stanleyi . In lateral profile, the cranium is distinctly straight, rising only gently up from the rostrum to the top of the braincase. An occipital ‘helmet’ is present but poorly developed, and the sagittal and lambdoidal crests are visible. The zygomatic arches are relatively robust ( Fig. 7 ), as in Lae. capensis . The dentition in Lae. kirinyaga is typical of the genus, with I 2/3, C 1/1, P 1/3, M 2/3. In the upper tooth row, I 1 is unicuspid and I 2 is small, not reaching halfway up the length 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: This species has been captured infrequently across the highlands of Kenya on both sides of the Rift Valley. It is present in wet tropical forest (e.g. Kakamega forest, with ~ 1900 mm of rainfall per annum), less mesic montane forest (Marsabit National Park ) and relatively dry savanna woodlands (e.g. Lolldaiga Hills conservancy ~ 600 mm), hence aridity per se does not seem to be an important variable in its distribution. However, it has been recorded only at elevations> 1000 m (current records are all between 1160 and 1700 m), and this might be an important limit in its geographical distribution. We also include two specimens (FMNH 233035, 233036) from Murchison Falls National Park, Uganda (1180 m above sea level) in this new species. Two specimens from Ethiopia (identified as ‘ Neoromicia somalica ’ by Benda et al., 2016) also group with Lae. kirinyaga in the phylogeny, as does a specimen from Senegal ( Koubínová et al., 2013), suggesting that this newly described species has a wide distribution north of the equator. We recommend, based on its relatively large distribution range and habitat preference, that it be listed as ‘Least Concern’ in the IUCN red list. However, we did not examine the specimens from Ethiopia and Senegal and therefore recommend a detailed morphological investigation before our hypothesis concerning the geographical range of this species is accepted. The type specimen echolocated at a peak frequency (start and end frequencies) of 44.9 kHz (74.3–41.6 kHz). The mean (± SD) peak frequency for 14 other individuals at the type locality was 43.9 ± 0.91 kHz (73.9 ± 9.43 to 41.8 ± 1.64 kHz).

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 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 6. Principal components analysis of craniodental characters of the short-eared species of Laephotis as recognized in this study.

Figure 7. Plate showing the cranium and mandible of Laephotis kirinyaga (FMNH 234558). Scale bar = 5 mm.

Figure 8. A, portrait of Laephotis kirinyaga (FMNH 234558), showing bright brown upper parts and off-white under parts, with bicoloured hairs. The skin around the eye is blackish in the holotype, but distinctly pinkish in most of the paratypes. B, portrait of Pseudoromicia nyanza (FMNH 215626), showing the distinctive white wings and under parts of this species.

Tables

Table 2. External measurements (in millimetres) and mass (in grams) of Laephotis kirinyaga from Marsabit National Park, Kenya

Specimen or taxon	Total length	Tail length	Hindfoot length	Ear length	Forearm length	Body mass
Laephotis kirinyaga	80	32	6	12	31.5	4.7
Holotype FMNH 234558
Laephotis kirinyaga	82.8 ± 5.67, 74–95,	32.6 ± 2.22, 28–37,	6.5 ± 0.97, 5–8,	11.5 ± 1.10, 9–13,	31.5 ± 1.35, 30.5–34.1,	5.3 ± 0.87, 4–7,
(other specimens)	N = 16	N = 16	N = 16	N = 16	N = 16	N = 16
Laephotis capensis	88.7 ± 7.09, 76–102,	34.9 ± 3.83, 26–40,	7.4 ± 0.84, 6–8,	12.8 ± 1.46, 11–17,	33.4 ± 2.22, 29.1–37.1,	5.7 ± 1.23, 3–8,
	N = 24	N = 24	N = 24	N = 24	N = 38	N = 18
Laephotis stanleyi	93.3 ± 6.42, 83–101,	39.7 ± 4.46, 34–48,	7.3 ± 0.76, 6–9,	12.3 ± 1.91, 10–15,	37.0 ± 1.58, 34–39,	6.5 ± 0.30,
	N = 8	N = 5	N = 8	N = 8	N = 8	6.1–6.8, N = 7
Laephotis robertsi	91.0 ± 4.06, 84–94,	34.6 ± 2.07, 31–36,	5.4 ± 0.55, 5–6,	13.0 ± 0, 13–13, N = 5	35.5 ± 1.41, 34.5–38.0,	8.5 ± 1.72,
	N = 5	N = 5	N = 5		N = 5	7.4–11.5, N = 5
Laephotis matroka	82.1 ± 3.18, 77–86,	32.2 ± 2.76, 27–36,	4.6 ± 0.51, 4–5,	11.8 ± 0.76, 11–13,	32.1 ± 1.51, 30–34,	5.3 ± 0.88,
	N = 12	N = 12	N = 12	N = 12	N = 12	4.0–7.5, N = 12
Laephotis malagasyensis	81.3 ± 1.15, 80–82,	36.0 ± 1.00, 35–37,	4.8 ± 0.50, 4–5,	12.0 ± 0.82, 11–13,	31.3 ± 0.96, 31–32,	4.5 ± 0.53,
	N = 3	N = 3	N = 4	N = 4	N = 4	3.7–6.0, N = 4

Measurements are presented as the mean ± SD, range and sample size (N). Measurements are of the holotype, other individuals of the new species and other ‘short-eared’ species of Laephotis . Measurements for the three Malagasy endemics, Lae. robertsi , Lae. matroka and Lae. malagasyensis , are taken from Goodman et al. (2017).