Murina alvarezi sp. nov.

(Figs. 1, 6, 7, 8E, 9E, 10A; Tables 1, 2A, 2B)

Murina cyclotis: Kitangland: Heaney et al. 1998: 27; Heaney et al. 2006: 30, 35, 36. Bohol Island: Sedlock et al. 2014: 205, 207; Siquijor Island: Heaney et al. 1998:27

Holotype. FMNH 205831, field number JLS 738, young adult female, specimen stored in alcohol, skull extracted, collected by J. L. Sedlock, 7 August 2009. The holotype is currently housed at FMNH but will be transferred to the National Museum of the Philippines, Manila before the end of 2025, where it has been assigned PNM 9670. The nucleotide sequence as deposited in GenBank of Cytb mitochondrial gene is PV659290.

Type locality. Sitio Bulalacao, Brgy Nug-as, Alcoy Municipality, Cebu Province, Cebu Island, Philippines, 9.7205 N 123.4594 E, elevation 763 m.

Paratypes. All seven paratypes were collected at the type locality: FMNH 205826, field number JLS 666, ♀, collected 25 July 2009; FMNH 205827, field number JLS 706, ♂, collected 3 August 2009; FMNH 205828, field number JLS 722, ♀, collected 3 August 2009; FMNH 205829, field number JLS 723, ♀, collected 3 August 2009; FMNH 205830, field number JLS 735, ♂, collected 3 August, 2009; FMNH 205832, field number JLS 739, ♀, collected 7 August 2009; and FMNH 205833, field number JLS 740, ♀, collected 7 August 2009 .

Etymology. This species is named in honor of James Alvarez, a remarkably devoted, young Filipino wildlife biologist with a passion for studying and conserving Philippine bats.A graduate student and biologist at the University of the Philippines —Los Baños Museum of Natural History, his unexpected death on Mount Apo in 2018 cut short a promising career of exploring Philippine biodiversity and inspiring future generations of Filipino biologists. The suggested English common name is Alvarez’s tube-nosed bat.

Measurements of holotype (in mm) and body mass (in g): total length, 97; tail vertebrae, 37; hind foot, 10.5; ear, 17, forearm, 35.6; and mass, 6.4.

Referred Material. FMNH 146890, field number SMG 5001, ♂, collected 21 February 1992, NW slope Mt. Guitingguitlin, 4.5 km S, 4 km E Magdiwang, Romblon Province, Sibuyan Island, Philippines, (12 27 N, 122 33 E, elevation 325 m) ; FMNH 202815, field number SEW 112, ♂, collected 11 July 2008 at Nan-od Brgy, Sierra Bullones Munic., Bohol Province, Bohol Island, Philippines (9.7469 N, 124.2638 E, elevation 586 m) ; FMNH 202816, field number JLS 590, ♂, collected 3 July 2008 at Sitio Libertad, Bugsoc Brgy, Sierra Bullones Munic. Bohol Province, Bohol Island, Philippines (9.7641 N, 124.2658 E, elevation 336 m) ; FMNH 209658, field number JLS 808, ♀, collected 6 km E of San Juan town centre, Siquijor Province, Siquijor Island, Philippines (9.1410 N 123.5472 E, elevation 170 m) ; FMNH 209659, field number JLS 809, ♂, collected 5 July 2010, same locality as previous; FMNH 209660, field number JLS 784, ♂, collected 23 June 2010 same locality as previous, 9.18706 N, 123.58040 E, elevation 550m.; FMNH147078, field number LRH 4639, ♂, collected 18 April 1992 Mt. Kitanglad Range, 115 km S, 6 km E Baungon, Bukidnon Province, Mindanao, Philippines (8 11 N, 124 44.5 E, elevation 1100 m) ; FMNH 147079, field number LRH 4665, ♀, collected 21 April 1992, same locality; FMNH 190119, RKSG 80, ♂, collected Hilong-hilong Range Carrasacal Munic., Suragio del Sur Prov. Mindanao; and FMNH 186817, field number DSB 3656, ♀, collected 19 May 2005 at Mt. Hamiguitan, San Isidro Munic., Davao Oriental Province, Mindanao, Philippines (6 43 56.3 N 126 09 03.2 E) .

Diagnosis. A medium sized Murina similar to Murina cyclotis (as defined by Francis & Eger 2012) but with a larger, heavier skull and genetically different at greater than 18% average sequence divergence in Cytb from other species in the genus (Table 3). Among the Philippine species of the M. cyclotis species group described here, M. alvarezi is the smallest (FA females = 35.8 mm, males = 33.7 mm; GLS females = 18.3, males = 17.5 mm); wing membrane attaches close to the claw of the toe; and penis narrow (1.5 mm). Similar to M. cyclotis, with all upper premolars similar in height; anterior upper molars with reduced mesostyles; and lower molars with reduced talonids (Figs. 8E, 9E). The sagittal and lambdoidal crests are clearly defined.

Description. In the holotype and paratype series of specimens from Cebu, the dorsal fur is bicoloured, grey buff at the base with reddish brown tips (Fig. 10A); the fur is 10–11 mm at the shoulder. On the underside, the throat is unicoloured, varying from white to buff; the ventrum is unicoloured white to buff in the centre and weakly bicoloured laterally, buff-grey to buff base and reddish brown to brown tips. The forearm, lower leg, interfemoral membrane, and foot are hairy. There is some colour variation within the species, with females from Siquijor and Mindanao paler than Cebu specimens. The ear is round and notched on the posterior edge (Fig. 10A). The skull has a moderately inflated braincase, shallow rostral depression, and small but defined sagittal and lambdoid crests. The upper toothrows are parallel to each other; the inner upper incisors are bicuspidate and longer than the outer; and the canines are large and heavy. The upper premolars are well developed, of similar height, and 2/3 the height of the canines (Fig. 8E). The mesostyles of the first (M 1) and second upper molars (M 2) are greatly reduced. The talonids (posterior section of the first, M 1 and second lower molars, M 2) are also greatly reduced relative to the trigonids (anterior section), about 1/3 the size of the trigonid (Fig. 9E). Penis of males is narrow (1.5 mm). See Tables 1 and 2A, 2B for selected external and skull measurements.

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Ecology. This species has a broad elevational range, occurring from 85 m elevation to 1100 m (Fig. 5) in a diversity of forest types and levels of disturbance. A single individual was captured in regenerating logged dipterocarp forest growing on ultramafic soils in southeastern Mindanao, characterized by stunted trees and an understory dominated by Malabayabas ( Tristaniopsis spp.) (Balete et al. 2006). Another individual was captured in northeastern Mindanao in a lowland Magkuno or ironwood ( Xanthostemon verdugonianus) forest. Several individuals were captured in second-growth dipterocarp forest on limestone in the Visayas, in small forest patches surrounded by corn and vegetable crops (Cebu, Siquijor and Bohol Islands; Sedlock et al. 2014); a mahogany plantation also on limestone; and within disturbed old-growth dipterocarp forest at 1100 m on Mt. Kitanglad (Heaney et al. 2006). Post-lactating females were captured in May in southeastern Mindanao (Mt. Hamiguitan), and in August on Cebu Island, along with three immature females. All individuals were captured in harp traps set along forest trails, except for two mist net captures on Mt. Kitanglad in central Mindanao, and one on Mt. Hamiguitan.

Distribution. Widespread within the southern and central Philippines, collected on Mindanao, Siquijor, Bohol, Cebu, and Sibuyan Islands (Fig. 1).

Genetic Analyses. Cytb and Dby sequences are available on GenBank for specimens listed in Appendices 2 and 3. Murina alvarezi is one of five divergent Cytb lineages from the Philippines that is related to M. cyclotis from mainland Asia and is supported by DBY data (one site, Fig. 2). Relative divergences among Philippine species are discussed below.