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Podogymnura Mearns, 1905 Type species: Podogymnura truei Mearns, 1905: 436. Included species: The type species, plus P. aureospinula, P. intermedia n. sp., and P. minima (raised to species rank, below). Distribution: Currently known from Bucas Grande Island, Dinagat Island, and the mountains of eastern (Mt. Hamiguitan and Kampalili) and central (Mts. Apo and Kitanglad) Mindanao Island (Fig. 1). Emended diagnosis: The genus Podogymnura is defined phylogenetically as the most recent common ancestor of P. aureospinula, P. truei, P. minima, and P. intermedia n. sp., and all of its descendants, and by the following combination of morphological characters. Small to medium-sized gymnures, HB from ca. 145 mm to 201 mm, CIL from ca. 39 to 52 mm (Tables 1 and 2); dorsal pelage reddish- or grayish-brown to dark brown with variable levels of golden highlights, soft to bristly and variable in length; underfur of woolly hairs fine, soft, and wavy; most guard hairs long, straight, and tapering with golden-yellow or golden-brown tips of varied lengths; black guard hairs long, distally flattened, tapered, and slightly bent. Two to three supragenal vibrissae present. Snout long, slender, and pointed, blunt at tip; nostrils extending slightly laterally, somewhat tubular; ear size moderate, extending far beyond pelage, sparsely covered on both surfaces with short, inconspicuous hairs; tail short relative to head and body, covered with short hairs that emerge posterior to the conspicuous scales that cover the tail; hind feet fairly long and narrow, covered thickly dorsally and thinly ventrally with short hairs. Females with two pairs of small, inconspicuous mammae, one pair inguinal and one pair axial. Adult males with slight swelling in uro-genital area, sometimes with bare skin from base of tail to area around penile sheath; no scrotum is evident. Rostrum elongate and broad, postorbital processes absent, frontals slightly to strongly inflated; interorbital region moderately to strongly constricted; braincase inflated; sagittal crest inconspicuous to prominent, extending no further anterior than anterior tips of interparietals, except extending further anterior then spreading into low temporal crests in P. aureospinula; nuchal crest of varying height that originates laterally as low projections from dorsal margin of each mastoid that meet mid-dorsally to form a broadly tapered arch with which the sagittal crest converges; incisive foramina cordate, moderately narrow, and short; anterior palatine foramina small and anterior to the maxilla/palatine suture; infraorbital canal dorsal or posterodorsal to the P4-M1 region; ante-orbital fossa present; posteroventral process of maxillary portion of zygoma present, small to prominent; ophthalmic foramen joined with or closely adjacent to the ethmoid foramen; lateral fossa absent from palatine anterodorsal to the post-palatal torus. Paired concavities in the basioccipital between the bullae, lateral to a medial ridge present in P. aureospinula and P. intermedia n. sp., absent in P. truei and P. minima. Upper molariform teeth with extreme anterior placement relative to orbit and infraorbital foramen; all incisors single-rooted; I2 and I3 similar in size, and both substantially smaller than I1; lower incisors spatulate and procumbent, i1 and i2 longer than i3; upper and lower canines double-rooted, flared laterally, and longest of all teeth. P1 and p1 absent; P2 single-rooted and small compared to other premolars, P3 larger antwo- or three-rooted, with lingual lobe small (P. aureospinula and P. intermedia n. sp.) or absent (all others); P4 large, broad lingually, and nearly square-shaped; M1 and M2 square-shaped, each with a low but discernable metaconule; metacone present on M3; c1 significantly larger than p1; lower molars sometimes with small cusp at base of talonid notch between entoconid and metaconid. Mandible relatively long and thick, with angular process narrow and long, coronoid wide, and condyloid process long and robust, especially in P. aureospinula., Published as part of Balete, Danilo S., Heaney, Lawrence R., Rickart, Eric A., Quidlat, Roselyn S., Rowsey, Dakota M. & Olson, Link E., 2023, A re-assessment of diversity among Philippine gymnures (Mammalia: Erinaceidae Podogymnura), with a new species from eastern Mindanao, pp. 244-266 in Zootaxa 5228 (3) on page 255, DOI: 10.11646/zootaxa.5228.3.2, http://zenodo.org/record/7532470, {"references":["Mearns, E. A. (1905) Descriptions of new genera and species of mammals from the Philippine Islands. Proceedings of the United States National Museum, 28, 425 - 460. https: // doi. org / 10.5479 / si. 00963801.1402.425"]}

Podogymnura minima Sanborn, 1953 Podogymnura truei minima Sanborn 1953. Mammals from Mindanao, Philippine Islands collected by the Danish Philippine Expedition, 1951-1952. Videnskabelige Meddelelser Dansk Naturhistorisk Forening 115:283–288. Podogymnura truei: Heaney & Morgan 1982. Tab. 1-2 (part, DMNH 5949-5953).— Poduschka & Poduschka 1985. Tab. 1 C-D, 2B, Abb. 6, 16 (part, FMNH 92777, 92780-81, SMF 31430, 31443, 31755, AMNH 164482; ZMC 1311).— Heaney et al. 2006. Tables 2 – 6, Fig. 8 (part, FMNH 74852, 92777, 92778, 92779, 92780, 92781,146592, 146594, 146962, 146963, 146964; 147782, 147783, 147790, 147800, 147801, 147802, 147803, 147804, 147808, 147809, 147810, 147811, 147812, 147813, 147814, 147815, 148003, 148004, 148005, 148006, 148010, 148011, 148012, 148013, 148016, 148017, 148018, 148019, 148050, 148053, 148077, 148078, 148083, 148084, 148085, 148087, 166456, 166457, 166458, 167375, 167376, 167377, 167378). Holotype. Zoologisk Museum, Copenhagen, catalog number 1311. Adult female collected on 16 December 1951, field number F. Salomonsen K-4. Skin and skull. Type locality. Mt. Kitanglad, Bukidnon Province, Mindanao Island, Philippines, 1600 m (Fig. 1). Measurements. Tables 1 and 2. Specimens examined. Mindanao Island, Bukidnon Province, Kitanglad Range, Mt. Kitanglad, 10.7 km S, 2.9 km W of Sumilao Poblacion, 1,450 m elev., 8°11’10”N, 124°55’10”E (FMNH 166456, 166457, 167375 – 167378); Mt. Kitanglad, 11.5 km S, 2.2 km W of Sumilao Poblacion, 1,500 m elev., 8°11’0”N, 124°55’35”E (FMNH 166458); Mt. Kitanglad, 1,600 m elev. (FMNH 74852); Mt. Imbayao, 15 km S, 7 km E of Baungon, San Vicente Municipality, 1,800 m elev., 8°9’N, 124°45’E (FMNH 146592, 146594, 146962 – 146964); Mt. Nangkabulos, 16.5 km S, 4 km E of Camp Phillips, 1,900 m elev., 8°10.5’N, 124°51’E (FMNH 147782, 147783, 147808, 147809, 147810, 147790, 148077, 148078); Mt. Nangkabulos, 15.5 km S, 4 km E of Camp Phillips, 2,250 m elev., 8°9.5’N, 124°51’E (FMNH 147800 – 147804, 147811, 147812, 147813, 147814, 147815, 148083, 148084, 148085, 148087); Mt. Dulangdulang, 15 km S, 11 km W of Dalwangan, Malaybalay City, 2,375 m elev., 8°7.5’N, 124°56’E (FMNH 148003, 148004, 148005, 148006, 148018, 148050); Mt. Dulang-dulang, 15 km S, 11.5 km W of Dalwangan, Malaybalay City, 2,600 m elev., 8°7.5’N, 124°56’E (FMNH 148010, 148011, 148012, 148013); Mt. Dulang-dulang, 15 km S, 12.5 km W of Dalwangan, Malaybalay City, 2,800 m elev., 8°7.5’N, 124°56’E (FMNH 148016, 148017, 148018, 148019, 148053); Mt. Kitanglad, Malaybalay City, 5,000 ft. elev. (ca. 1,524m), (FMNH 92777, 92778, 92779); Mt. Kitanglad, Malaybalay City 6,000 ft. elev. (ca. 1,829 m), FMNH 92780, 92781). Distribution. Documented from Mt. Kitanglad Range, north-central Mindanao Island, including Mts. Dulangdulang, Imbayao, and Nangkabulos (Fig. 1). Emended diagnosis. Overall, the smallest species of the genus (HB = 137 – 152 mm); tail short (49 – 66 mm), about 36% of head and body, pale grayish-brown dorsally and unpigmented ventrally; hindfoot short (HF = 33 – 37 mm, 23% of HB; Table 1) and uniformly pale brown. Dorsal pelage dark reddish-brown, long and soft with conspicuous golden-brown tips; underfur dark gray, dense and wavy, shorter than guard hairs; guard hairs black, with most long, straight, and tapered, but some distally flattened and slightly bent. Ears and feet pale, lightly pigmented. Skull (Fig. 5, Table 2) slender and tapered (CIL = 37.28 – 39.78 mm), sagittal and nuchal crests poorly developed and inconspicuous, rostrum long (LR = 15.00 – 17.57 mm), cranium narrow (BBC = 15.40 – 16.22 mm). Incisive foramina relatively narrow and short. Anterior surface of basioccipital nearly smooth, lacking or barely showing a short ridge running medially parallel to the bullae; paired concavities absent. Tips of tympanic wings of basioccipital short and nearly straight (rather than longer and curved medially). Upper toothrow short (I1 – M3 = 18.71 – 21.06 mm), P3 without lingual lobe; mandible relatively slender (LMI = 28.63 – 30.65 mm), as are its angular, coronoid and condyloid processes. P4 relatively small and triangular. Cusp at base of talonid on m1 and m2 absent. Comparisons. Podogymnura minima and P. aureospinula: P. minima is the smallest and P. aureospinula the largest member of the genus (Tables 1 and 2), and they are easily distinguished on that basis. The long, soft pelage of P. minima strongly contrasts with the short and stiff, bristly fur of P. aureospinula; in the former species, goldenbrown highlights visible only at the tips of black guard hairs, but the latter has conspicuous golden-yellow guard hairs. Paired concavities in basioccipital between bullae that are present in P. aureospinula and P. intermedia are absent. The lingual lobe of P3 is present in P. aureospinula but absent in P. minima. P4 relatively small and triangular, vs. large and squarish. Cusp at base of talonid on m 1 and m 2 absent, vs. present. Podogymnura minima and P. intermedia n. sp: pelage of P. minima is soft, similar to that of P. intermedia from Mt. Kampalili, and different from the rough and bristly dorsal pelage of P. intermedia from Mt Hamiguitan. P. intermedia has conspicuous dorsal golden-yellow streaks or speckling, whereas P. minima has smaller, less apparent golden-brown speckles. P. minima is smaller than P. intermedia n. sp. in nearly all respects, especially the large specimens from Mt. Hamiguitan (Tables 1 and 2, Figs. 6A and 6B). It is notable that P. intermedia from Mt. Hamiguitan has an especially long and broad rostrum, long post-palatal region, and thick mandible relative to P. minima (Table 2). Further comparisons in the new species description below. Podogymnura minima and P. truei: See comments above; external differences include small overall size in P. minima (mean HB = 145 mm vs. 148 mm, TV = 54 vs. 56 mm). The skull of P. minima is slightly but consistently shorter and more gracile overall, with a narrower rostrum, less inflated braincase, and lower sagittal crest; P. truei has slightly shorter condylar process of the mandible (Tables 2, 3, Figs. 5, 6A, 6B). Karyology. Specimens from the Kitanglad Range have a standard karyotype of 2N = 40, FN = 76 (Rickart 2003). Ecology. P. minima has been recorded in montane and mossy forest, from 1300 m to 2800 m elevation; it was among the most abundant small mammals in middle to high-elevation montane and mossy forest on Mt. Kitanglad (Heaney et al. 2006). They are nocturnal, feeding on the surface of the ground. Diet based on stomach contents is composed largely of earthworms, with some arthropods, including hymenopterans and coleopterans. Pregnancy was recorded from March to June; litter size (n = 9) was one, rarely two. Other species of native small mammals documented in the elevational range of P. minima were Crocidura beatus, Tupaia everetti, Apomys hylocoetes, A. insignis, Batomys salomonseni, Crunomys suncoides, Limnomys bryophilus, L. sibuanus, Rattus everetti, and Tarsomys apoensis (Heaney et al., 2006)., Published as part of Balete, Danilo S., Heaney, Lawrence R., Rickart, Eric A., Quidlat, Roselyn S., Rowsey, Dakota M. & Olson, Link E., 2023, A re-assessment of diversity among Philippine gymnures (Mammalia: Erinaceidae Podogymnura), with a new species from eastern Mindanao, pp. 244-266 in Zootaxa 5228 (3) on pages 256-257, DOI: 10.11646/zootaxa.5228.3.2, http://zenodo.org/record/7532470, {"references":["Sanborn, C. C. (1953) Mammals from Mindanao, Philippine Islands collected by the Danish Philippine Expedition, 1951 - 1952. Videnskabelige Meddelelser Dansk Naturhistorisk Forening, 115, 283 - 288.","Heaney, L. R. & Morgan, G. S. (1982) A new species of gynmure, Podogymnura, (Mammalia: Erinaceidae) from Dinagat Island, Philippines. Proceedings of the Biological Society of Washington, 95, 13 - 26.","Poduschka, V. W. & Poduschka, C. (1985) Beitrage zur Kenntnis der Gattung Podogymnura Mearns 1905 (Insectivora: Echinosoricinae). Zeitschrift fur Saugetierkunde, 50, 1 - 21.","Heaney, L. R., Tabaranza, B. R. Jr., Rickart, E. A., Balete, D. S. & Ingle, N. R. (2006) The mammals of Mt. Kitanglad Nature Park, Mindanao, Philippines. Fieldiana: Zoology (New Series), 112, 1 - 63. https: // doi. org / 10.3158 / 0015 - 0754 (2006) 186 [1: TMOMKN] 2.0. CO; 2","Rickart, E. A. (2003) Chromosomes of Philippine mammals (Insectivora, Dermoptera, Primates, Rodentia, Carnivora). Proceedings of the Biological Society of Washington, 116, 473 - 487."]}

Podogymnura intermedia new species Figs. 5, 7, 8, and 9; Tables 1 and 2 Holotype. FMNH 186805, adult male, collected on 28 July 2005 (original number 3769 of D. S. Balete); initially fixed in formalin, now preserved in 70% ethyl alcohol with the skull removed; skull in good condition. A sample of fresh muscle tissue was removed from the left thigh and preserved in 90% ethanol; otherwise the body is in good condition. The holotype has been cataloged and is currently housed at FMNH but will be transferred to the National Museum of the Philippines, Manila. Type Locality. 3.7 km S, 1.4 km E of Mt. Hamiguitan peak, Mati Municipality, Davao Oriental Province, Mindanao Island, Philippines, 950 m elev., 6 º 42’ 26.2” N, 126 º 11’ 42.8” E (Fig. 1). Specimens examined. Mt. Hamiguitan: (N = 4) Three additional specimens from the type locality (FMNH 190152, 190153, 190167) and one (FMNH 190151) from 17.5 km S, 4 km E of Mt. Hamiguitan peak, San Isidro Municipality, ca. 1,128 m elevation, 6 º 43’3” N, 126 º 11’1.9” E. These include two males and one female, and a specimen of undetermined sex prepared as a skull only (FMNH 190167). The juvenile male (FMNH 190151), adult male (FMNH 190152), and young adult female (FMNH 190153) have had their skulls removed and cleaned. The bodies of the adult male and young-adult female had partially decomposed prior to preservation, but that of the juvenile male is in good condition. The prepared skulls have broken crania, including one with broken coronoid and condyloid processes (FMNH 190152). Mt. Kampalili: (N = 12) Three males (FMNH 194750, 194751, 194752) from 2 km S, 2 km W of Mt. Kampalili peak, Maragusan Municipality, Compostela Valley Province, Mindanao Island, Philippines, ca. 1,900 m elevation, 7° 17’ 11.7” N, 126° 15’ 30.9” E; two females (FMNH 194748, 194749) from 2.75 km S, 0.5 km W of Mt. Kampalili peak, ca. 1,500 m elevation, 6 º 17’39” N, 126 º 15’38.4” E; two females (FMNH 208700, 208701) and three males (FMNH 208699, 208702, 208705) at 2.25 km S, 3.5 km E of Mt. Kampalili peak, 1,470 m elevation, 7.29112 º N, 126.31520 º E; and two females (FMNH 208703, 208704) at 1.75 km S, 4.25 km E of Mt. Kampalili peak, 1,640 m elevation, 7.29522 º N, 126.31602 º E. Three females (FMNH 194748, 208700, 208703) and three males (FMNH 194750, 194752, 208705) have had their skulls removed and cleaned; others preserved intact. Two females (FMNH 194748, 208700) have crushed crania, and one male (FMNH 194752) has a broken left zygomatic arch; and one male (FMNH 208705) has a short crack on the occipital. Distribution. Currently known from montane forest over ultramafic soil between ca. 950 and 1,128 m on Mount Hamiguitan, and in montane and mossy forest from ca. 1,470 m to 1,900 m on Mount Kampalili (Fig. 1). Etymology. From the combined Latin inter (between) + medius (middle), to highlight its intermediate morphology between the two smaller species of Podogymnura and the large P. aureospinula. We propose eastern Mindanao gymnure as its English common name. Diagnosis. A member of the genus Podogymnura as diagnosed above, of intermediate to small size (Mt. Hamiguitan average HB = 171 mm, weight 96 g; Mt. Kampalili average HB = 145, weight 77 g; Table 1), defined by the following combination of characters. Dorsal pelage generally coarse in Hamiguitan, softer in Kamapalili specimens, with long guard hairs of two types: scattered long, black hairs, and many slightly shorter guard hairs having conspicuous golden-yellow tips that produce an appearance of shiny, golden streaks (Fig. 7). Both types of guard hairs become progressively shorter laterally and anteriorly over the head and snout. Underfur soft, gray, and dense, becoming sparser over head but more conspicuous laterally as guard hairs become sparser laterally and absent ventrally, where silver highlights become apparent. Ventral pelage shorter and paler, lacking golden streaks. Ears short (average = 20 mm), pigmented grayish-brown. Relative to congeners, hindfoot longer in Hamiguitan (average HF = 36 mm), shorter in Kampalili (34 mm) but short relative to HB (21–23 % of HB; Table 1); plantar surface, including plantar pads, darkly pigmented or with mottled grayish-brown areas. Tail pigmented medium to dark grayish brown dorsally, and variable ventrally, usually dark but occasionally mottled with white. Tail relatively short compared to length of head and body (average TV = 53–54 mm; 32–37 % of HB; Table 1). Skull of Hamiguitan specimens (Figs. 5, 8) large (holotype CIL = 43.4 mm, BBC = 16.8 mm), that of Kampalili specimens smaller, (CIL = 40.6 mm, BBC = 16.1 mm). Sagittal and nuchal crests low but readily visible, rostrum long and deep. Incisive foramina relatively long and wide in Hamiguitan specimens, narrower and shorter in Kampalili specimens. Upper toothrow long (average I1– M3 = 21.1 – 22.2 mm). Post-palatal region proportionately long in specimens from Mt. Hamiguitan, average in Kampalili, both samples with paired concavities between the bullae, bisected by a low medial ridge. Mandible long and robust in Hamiguitan (average LMI = 33.5 mm), shorter and more slender in Kampalili (average LMI = 31.6 mm). Lingual lobe of P3 present in Hamiguitan specimens, absent in Kampalili. P4 proportionately large and square-shaped. A small, inconspicuous cusp present at base of talonid on first and second lower molars (Fig. 8E). Description and Comparisons. Because specimens from Mt. Hamiguitan (the type locality) differ from those from Mt. Kampalili, we include comparisons of specimens from these two places along with comparison to other species of Podogymnura. A medium-sized Philippine gymnure, Hamiguitan specimens larger and more robust than specimens from Mt. Kampalili, P. minima, and P. truei, but smaller and less robust than P. aureospinula (Tables 1 and 2, Figs. 4, 5). Dorsal pelage of P. intermedia from Hamiguitan is distinctly darker on head and rostrum, venter slightly paler brownishgray (Fig. 4); coarse overfur consists of long, stiff guard hairs, ca. 20 mm on mid-dorsum, longer on the rump, ca. 25 mm. The long, coarse, grayish-brown dorsal pelage contains both long black guard hairs and long golden-yellow guard hairs that produce a golden-streaked appearance. Mt. Kampalili specimens have soft, dark brown dorsal pelage with shorter black guard hairs and many short golden-yellow-tipped hairs that produce a golden-speckled appearance. Dorsal pelage of P. aureospinula is uniformly paler and golden-brown overall and overfur is shorter and bristly; venter brownish-gray. P. truei has soft dorsal pelage that is dark brown with small, inconspicuous speckles of golden-brown at the tips, without conspicuous long and stiff guard hairs. P. minima has short, soft, reddish-brown dorsal pelage with golden-brown speckles at the tips that are more conspicuous than those of P. truei but are darker and smaller than those of P. intermedia from Mt. Hamiguitan or specimens from Mt. Kampalili. Females of all species have two pairs of small, inconspicuous mammae, one pair inguinal and one pair axial; these are well hidden by the fur.Adult males have a swollen area from the base of the tail to the area around the penile sheath; the sheath is small, ca. 1.5 mm wide and long; no scrotum is evident. In specimens from Hamiguitan, the swollen area is covered by typical abdominal fur, but in specimens from Kampalili, the swollen area is bare. Ears of all Podogymnura are short relative to body size, and sparsely covered with short, nearly invisible hairs; ears of specimens from Hamiguitan are pigmented dark gray but from Kampalili are paler. P. aureospinula ears are palest of the known species. Ears of P. truei and P. minima average slightly longer and paler than in P. intermedia (Table 1). Length of tail relative to head and body (32%) in P. intermedia from Hamiguitan is shortest among all Podogymnura (Table 1). Skin of tail of Hamiguitan specimens is uniformly dark gray throughout; on Kampalili specimens it is ventrally pale brown or mottled with white. Tail of P. aureospinula is longer but overlaps in relative length (33% of HB). Tail of P. truei is substantially longer both absolutely and proportionately (Table 1). Although P. minima is smaller overall, tail is equal in length and proportionately longer. Hind foot of P. intermedia is average among Podogymnura excepting the larger P. aureospinula, but proportionately shorter than all except P. aureospinula (Table 1). Hindfoot skin is pigmented medium gray dorsally and ventrally, including digits and plantar pads, paler and often mottled with white in those from Kampalili. Hind feet of P. aureospinula, P. truei and P. minima are unpigmented, both dorsally and ventrally. Skull of P. intermedia (Fig. 8, Table 2) is smaller than the much larger P. aureospinula (Fig. 4), but larger than its congeners in most respects, with Mt. Hamiguitan specimens larger than those from Kampalili (Fig. 5). It is similar to P. truei and P. mimima in its limited development of the prominent nuchal and sagittal crests that are hallmarks of P. aureospinula. Maximum height of sagittal crest is 0.8 mm in P. intermedia. The nuchal crest of P. intermedia slants slightly posteriad relative to the cranium but does not project beyond occipital margins (Fig. 8), which it does in P. aureospinula (Fig. 4). In lateral view, skull of P. intermedia from Hamiguitan (Fig. 8) cuts a nearly straight slanting profile from top of braincase to tip of narrow and tapered rostrum, whereas in specimens from Kampalili the profile is slightly concave. Frontals of P. aureospinula are dorsolaterally inflated, making them only about 9% narrower than braincase, and in dorsal view producing a narrow-waisted hour-glass shape in interorbital region (Fig. 4). In lateral view, prominent frontal swellings in P. aureospinula produce a convex dorsal profile. All three small-bodied Podogymnura share an interorbital region in which there is a relatively broadly-waisted hourglass shape, although lacrimal and interorbital breadths of P. intermedia from Hamiguitan are greater than in Kampalili and the other two species (Table 2), and all lack the temporal ridges that in P. aureospinula form a low crest converging at the interorbital region. In P. intermedia and the two other small-bodied species, the parietals are dorsolaterally inflated from the anterior edge of interparietals, forming a cranium that tapers anteriad to the frontal region and flattens posteriad to the occipital region (Fig. 8). Incisive foramina cordate, similar to those of P. minima and P. truei in size, in contrast to the wider and longer foramina in P. aureospinula and P. intermedia. Palatal length, width of the posterior palatal ridge, and lingual palatal breadth at M 3 of P. intermedia are large (except in comparison to P. aureospinula; Table 2). Postpalatal length of P. intermedia from Hamiguitan is the greatest among the small Podogymnura, with specimens from Kampalili averaging shortest (Table 2). The basicranium of P. intermedia is similar to its congeners, differing mainly in the large size of the basicranial area, auditory bulla, and paraoccipital process (Fig. 8). Aside from overall size, the basicranium of P. intermedia is similar to that of P. aureospinula (and unlike other Podogymnura species) in having paired, shallow concavities traversed medially by a fine, short ridge that runs parallel to the bullae from its base to the tip of the tympanic wing, although in P. aureospinula the depressions are more expansive and the mid-ventral ridge longer and larger. The medial ridge is low and poorly defined in P. truei and nearly absent in P. minima, and the concavities are absent in both. Mandible of P. intermedia (Fig. 8, Table 2) relatively long and thick, except in comparison to P. aureospinula (Fig. 5). P. aureospinula has larger, wider, and longer angular, coronoid, and condyloid processes than all of its congeners. In P. intermedia from Hamiguitan, the angular process is narrower but longer, coronoid wider, and condyloid process longer and more robust than in specimens from Kampalili, P. minima, and P. truei. Position of mandibular foramina is similar in all species. Dental features of P. intermedia (Figs. 8, 9A, 9B, Table 2) are similar to those of congeners in most respects (see Podogymnura Diagnosis), usually differing only in relative sizes. The presence of a discernible lingual lobe in P 3 in specimens from Hamiguitan is shared with P. aureospinula; this lobe is absent among all other Podogymnura. P4 is proportionately larger and more square-shaped in P. aureospinula and P. intermedia than in P. truei or P. minima. A small, inconspicuous cusp at base of talonid on m1 and m2 is present in all specimens of P. intermedia from Hamiguitan (Fig. 8E) and P. aureospinula (Heaney & Morgan 1982); a poorly-developed version is present in a small percentage of P. truei, and is absent in other populations, including P. intermedia from Kampalili. Ecology. We recorded P.intermedia on Mt. Hamiguitan and Mt. Kampalili in different types of forest formations, and so we present the ecological information separately. On Mt. Hamiguitan, we captured P. intermedia in primary montane forest over ultramafic soil at 950 m to 1,128 m (Balete et al. 2006). We did not record it in lowland forest at 525 m in 924 trap-nights. At 950 m, three individuals were caught in 228 trap-nights with earthworm bait (1.3% success) compared to one individual caught in 672 trap-nights with roasted coconut coated with peanut butter (0.15%); none were captured in traps set above ground on vines and trees. On Mt. Kampalili, we recorded this species in upper montane and mossy forest at ca. 1,470 m – 1,900 m elevation; forested habitats below and above these elevations were not surveyed. Seven P. intermedia recorded during 2010 were captured in 247 trap-nights with live earthworm bait (2.83 % trap success); none were captured in 506 trap-nights using coconut bait. All were captured on the ground, and none in traps set in trees or on vines, and nearly all were captured at night. On the adjacent peak of Mt. Kangayag (ca. 1,630 m) of the same mountain range, we captured no gymnures in 2,140 trap-nights, of which 250 were baited with live earthworms. The apparent absence of P. intermedia in the lowlands and restriction to montane and mossy forest is similar to P. minima, which occurs in secondary and old-growth montane and mossy forest from ca. 1,300 m to 2,800 m (Heaney et al. 1998, 2006), and to P. truei, which has been recorded from montane and mossy forest at ca. 1,640 m to at least 2,250 m (Hoogstraal 1951, Sanborn 1952). In contrast, P. aureospinula occurs in secondary and oldgrowth lowland forest at low elevations (Heaney & Morgan 1982; Heaney & Rabor 1982; Heaney et al. 2010). Stomach contents of two individuals from Mt. Hamiguitan contained mainly chewed remains of arthropod exoskeletons, including coleopterans and their larvae, as well as an operculum and shell fragments of small land snails. A few pieces that appeared to be partially digested bits of small earthworms were present. Stomach contents of three specimens from Mt. Kampalili contained chewed remains of arthropod exoskeletons, including coleopterans and centipedes, and two contained pieces of earthworms. The presence of land snails in the diet of this species is the first record of this food item among Podogymnura. These diet and trapping data indicate that, as with other Podogymnura, P. intermedia forages on the ground for leaf-litter invertebrates. These feeding habits may differ from those of P. minima, which feeds more commonly on earthworms than arthropods (Heaney et al. 2006), although this difference may simply reflect differences in prey availability. An adult male from Mt. Hamiguitan taken in May had testes measuring 11 x 5 mm; an adult female was nulliparous. The presence of a juvenile in May suggests that some breeding occurred earlier in the year. None of the specimens from Mt. Kampalili captured in February and May (five adult males, three adult females, three young adult females and one young adult female) showed signs of reproductive activity. Three species of murid rodents (Batomys hamiguitan, Bullimus bagobus, and Rattus everetti) were documented as being sympatric with P. intermedia on Mt. Hamiguitan along with the Mindanao shrew (Crocidura beatus; Balete et al. 2006, 2008). In addition to these native species, we recorded the non-native spiny ricefield rat (Rattus exulans) at two disturbed sites near the type locality: a narrow patch of non-native cogon grass, Imperata cylindrica, surrounding the shoreline and dry lakebed of Tinagong Dagat, a seasonal lake; and in disturbed vegetation along a foot-trail, but not in the nearby forest interior. On Mt. Kampalili this species was associated with Apomys sp., Bullimus bagobus, Rattus everetti, Baletemys kampalili (Rowsey et al. 2022), an undescribed species of Tarsomys, and one squirrel, Sundasciurus philippinensis (specimens in FMNH). The commensal house shrew, Suncus murinus, was also present. All of the native co-occurring species are endemic to the Philippines, and all but R. everetti are restricted to the Mindanao Faunal Region (Heaney 1986; Heaney et al. 2006, 2010)., Published as part of Balete, Danilo S., Heaney, Lawrence R., Rickart, Eric A., Quidlat, Roselyn S., Rowsey, Dakota M. & Olson, Link E., 2023, A re-assessment of diversity among Philippine gymnures (Mammalia: Erinaceidae Podogymnura), with a new species from eastern Mindanao, pp. 244-266 in Zootaxa 5228 (3) on pages 258-262, DOI: 10.11646/zootaxa.5228.3.2, http://zenodo.org/record/7532470, {"references":["Heaney, L. R. & Morgan, G. S. (1982) A new species of gynmure, Podogymnura, (Mammalia: Erinaceidae) from Dinagat Island, Philippines. Proceedings of the Biological Society of Washington, 95, 13 - 26.","Balete, D. S., Quidlat, R. S. & Ibanez, J. C. (2006) The non-volant small mammals of Mt. Hamiguitan, Eastern Mindanao, Philippines. Banwa, 3, 65 - 80.","Heaney, L. R., Balete, D. S., Dolar, L., Alcala, A. C., Dans, A., Gonzales, P. C., Ingle, N. R., Lepiten, M., Oliver, W., Ong, P., Rickart, E. A., Tabaranza, B. R. Jr. & Utzurrum, R. C. B. (1998) A synopsis of the mammalian fauna of the Philippine Islands. Fieldiana: Zoology, New Series, 88, 1 - 61.","Hoogstraal, H. (1951) Philippine zoological expedition, 1946 - 1947. Narrative and itinerary. Fieldiana: Zoology, 33, 1 - 33.","Sanborn, C. C. (1952) Philippine zoological expedition, 1946 - 1947: Mammals. Fieldiana: Zoology, 33, 1 - 158.","Heaney, L. R. & Rabor, D. S. (1982) An annotated checklist of the mammals of Dinagat and Siargao islands, Philippines. Occasional Papers of the Museum of Zoology, University of Michigan, 699, 1 - 30.","Heaney, L. R., Dolar, M. L., Balete, D. S., Esselstyn, J. A., Rickart, E. A. & Sedlock, J. L. (2010) Synopsis of Philippine Mammals. Field Museum website, http: // www. fieldmuseum. org / philippine _ mammals /","Heaney, L. R., Tabaranza, B. R. Jr., Rickart, E. A., Balete, D. S. & Ingle, N. R. (2006) The mammals of Mt. Kitanglad Nature Park, Mindanao, Philippines. Fieldiana: Zoology (New Series), 112, 1 - 63. https: // doi. org / 10.3158 / 0015 - 0754 (2006) 186 [1: TMOMKN] 2.0. CO; 2","Balete, D. S., Heaney, L. R., Rickart, E. A., Quidlat, R. S. & Ibanez, J. C. (2008) A new species of Batomys (Muridae: Murinae) from eastern Mindanao Island, Philippines. Proceedings of the Biological Society of Washington, 121, 411 - 428. https: // doi. org / 10.2988 / 07 - 47.1","Rowsey, D. M., Duya, M. R. M., Ibanez, J. C., Jansa, S. A., Rickart, E. A. & Heaney, L. R. (2022) A new genus and species of shrewlike mouse (Rodentia: Muridae) from a new center of endemism in eastern Mindanao, Philippines. Journal of Mammalogy, 103. https: // doi. org / 10.1093 / jmammal / gyac 057.","Heaney, L. R. (1986) Biogeography of mammals in Southeast Asia: estimates of rates of colonization, extinction, and speciation. Biological Journal of the Linnean Society, 28, 127 - 165. https: // doi. org / 10.1111 / j. 1095 - 8312.1986. tb 01752. x"]}

Kerivoula pellucida Waterhouse 1854 Type locality: Philippines. Specimens examined. Given in Appendix A. Distribution. Borneo, Java, the Malay Peninsula, the Philippines (Cebu, Jolo, Mindanao, Mindoro, and Palawan), and Siberut (Fig. 2; see also Corbet & Hill 1992: 154). Description. Total length 83–97 mm, tail 43–50 mm, ear 16–18.5 mm, forearm 31.1–35.2 mm, tibia 16.6–19.4, mass 4.0– 5.7 mm (Table 4). Dorsal pelage is long and soft with a slightly woolly appearance, and pale reddish- brown buff; hairs slightly paler at base than at tips, lacking distinct color bands; ventral pelage paler (Fig. 5D, E). The ears are pale brown, and unusually large (Fig. 5A, B). A thin scattering of inconspicuous hairs is present over the dorsal surface of the interfemoral membrane, some of which project over its posterior edge (Fig. 5A). The wing and tail membranes are very pale brown, nearly translucent (Fig. 5A, C). Tragus long and slender (Fig. 5B). Skin is nearly transparent giving the face, wing- and leg bones a reddish-pink hue in live animals. The wing membrane inserts onto the side of the outer toe well above the base, not at its base (Fig. 5C). There do not appear to be striking regional differences within the Philippines in appearance; however, specimens from Bohol are slightly darker brown than those from Mindanao and Cebu; those from Palawan are palest. Palawan specimens have more reddish-brown tips on the dorsum than those from Bohol and Cebu. Skull as shown in Figure 6. GTL14.06–15.03, CCL 12.75–13.44 mm, GBB 7.04–7.55 mm, BH 5.69–6.33 mm, PC 2.98–3.15, C–M 3 5.63–6.05 mm, M 2 –M 2ext 4.88–5.16 mm, C 1 –C 1 1.62–1.86 mm, C–M 3 6.05–6.50 mm, MDL 10.19–11.16 mm (Table 1). The skull is narrow with a highly inflated braincase that forms a steep forehead. The post-palatal region extends far posterior to the molar tooth row, and the bullar cochleae are moderately large. The upper tooth rows are slightly convergent anteriorly. Infraorbital canal is short, above P 3 to middle of M 1. The upper incisors are conical; the first upper incisor is more than twice the height of the second upper incisor, which is often separated from the canine by a narrow space. The first lower incisor (Fig. 7B) usually has four cusps (occasionally three), the second has three cusps, and the third has two prominent cusps and usually a third (at the posterior edge). The upper and lower canines are high and conical with a cingulum that extends from the anterior edge lingually to the posterior margin, about twice the height of the first premolars. The first upper premolar is nearly circular in outline, and the second is slightly compressed laterally (Fig. 7A). The third upper premolar is partially molariform, with a broad lingual shelf. The first lower premolar is roughly circular in outline only slightly compressed laterally; the second and third are more compressed laterally, but not strongly so. The upper and lower molars are typical for the genus (Hill 1965). Philippine specimens are slightly larger than a small series from Malaysia (Fig. 3; Tables 1, 4). Comparisons. Kerivoula pellucida differs from K. whiteheadi in having dorsal and ventral pelage that is paler at the base than at the tips (Fig. 5D, E) (rather than dark at the base), reddish-brown at the tips, and slightly paler at mid-shaft, giving a tri-banded appearance. The wing and interfemoral membranes of K. pellucida are pale and nearly transparent, rather than moderately dark brown as those of K. whiteheadi. The tail membrane of K. pellucida has shorter, less conspicuous hairs (though still few and scattered) than that of K. whiteheadi. The wing membrane inserts up onto the side of the toe (Fig. 5C), rather than at the base of the outer toe. Most individuals of K. pellucida have greater total length than in K. whiteheadi, but much of this is due to having a longer tail (at least 43 mm, rather than 41 mm or less). The forearm is typically longer (usually 32 or more, rather than 31 mm or less), and the ear averages longer, though there is overlap (Table 4). The rostrum, palate, and braincase of K. pellucida are proportionately broader than those of K. whiteheadi (Fig. 6), the upper and lower premolars are less laterally compressed, there is more space between the molars lingually in K. pellucida, and the toothrows are typically longer (Fig. 7). The posterior palatal extension in K. pellucida is longer and narrower. Kerivoula pellucida differs from the Philippine K. hardwickii (i.e., K. “ hardwickii A” and K. “ hardwickii B”) in having larger and more pointed ears (rather than smaller and rounded; Heaney et al. 2010, 2016). The pelage of K. pellucida is paler at the base of each hair (Fig. 5D, E), rather than much darker at the base. Wing membranes of K. pellucida are pale and nearly transparent (Fig. 5A), rather than moderately dark brown and not transparent. Skin over the wing bones on individuals in the K. hardwickii group are darkly pigmented, not nearly transparent giving the bones the appearance of white stripes in preserved specimens and a pinkish hue in live animals, as in K. pellucida. Hairs on the tail membrane of Philippine K. hardwickii are very sparse and short, but a fringe of short hairs is often visible along the posterior edge. The wing membranes attach at the base of the outer toe, not part way up as in K. pellucida (Fig. 5C). The average total length of K. pellucida (91.4 mm) is longer than K. “ hardwickii A” (82.8 mm) and K. “ hardwickii B” (85.0 mm); but the average forearm length of K. pellucida (32.9 mm) is similar to K. “ hardwickii A” (33.0 mm) and smaller than K. “ hardwickii B” (35.0 mm) (Table 4; Heaney et al. 2010; unpublished data). The skull in K. pellucida is generally proportionately longer and narrower than Philippine K. hardwickii (Fig. 8). The post-palatal extension of Philippine K. hardwickii is broader and shorter than that of K. pellucida. Finally, the palate is proportionately narrower in K. pellucida than in Philippine K. hardwickii. Kerivoula pellucida differs from Philippine K. papillosa in being much smaller in all respects (Fig. 3; Heaney et al. 2010, 2016); the average forearm length of K. pellucida is 32.9 mm compared to 42.6 mm in K. papillosa (Heaney et al. 2010; unpublished data). The pelage of K. pellucida is much paler overall than K. papillosa, which is a dark brown. The skull is smaller and more delicate, with conspicuously narrower rostrum, palate, and braincase (Figs. 6, 9). The second upper incisor reaches only one-third toward the tip of the first incisor, rather than more than halfway (Figs. 7A, B, 10A, B). All of the premolars and molars are proportionately more massive. Echolocation. Philippine K. pellucida (4 individuals from Bohol, 1 from Cebu) have extremely broadband (132 kHz), short duration (2 ms), high frequency (peak = 138 kHz) calls similar to those recorded in Peninsular Malaysia (Fig. 11; Table 5; Kingston et al. 1999, Schmieder et al. 2010). They are also similar to those of Philippine K. whiteheadi with respect to peak frequency, although K. pellucida calls had a broader bandwidth (Table 5). Both species’ calls start at a similarly high frequency (194 kHz), but K. pellucida calls terminate at a lower frequency. Ecology. In the Philippines, K. pellucida occupies a broad range of elevations (sea level to about 1200 m), though most records are from below 700 m. They also occupy a wide range of habitats; examples include an isolated patch of second growth forest surrounded by vegetable farms on Cebu Island, disturbed second growth forest on limestone on Bohol Island, and pristine montane forest on Mindanao Island. Its roosting habits are poorly known in the Philippines, but one adult female with a suckling young was captured by hand from dried banana leaves on Palawan Island (J. Esselstyn, pers. com.). Payne et al. (1985) also reported it roosting in dried banana leaves, and Kingston (2006) reported a group roosting in a clutter of dead understory leaves. In Peninsular Malaysia, they roost in small groups of up to 15 individuals and appear to have small home ranges, evidenced by short recapture distances (et al. 2012). On Bohol Island, on three occasions, multiple individuals were captured either in the same or adjacent harp traps simultaneously, usually a male and a female (Sedlock et al. 2014). In Peninsular Malaysia, K. pellucida exhibits asynchronous reproduction giving birth throughout the year, although the highest proportion of pregnant individuals was captured during the rainy season when insect abundance was highest (Nurul-Ain et al. 2017). In the Philippines, five lactating females were captured in July, during the rainy season (Sedlock et al. 2014), and one female was captured with a suckling young in late March (J. Esselstyn, pers. com.). Its echolocation behavior is highly adapted for distinguishing prey from background clutter at close range allowing it to forage within the forest understory (Schmieder et al. 2012)., Published as part of Sedlock, Jodi L., Heaney, Lawrence R., Balete, Danilo S. & Ruedi, Manuel, 2020, Philippine bats of the genus Kerivoula (Chiroptera: Vespertilionidae): Overview and assessment of variation in K. pellucida and K. whiteheadi, pp. 454-490 in Zootaxa 4755 (3) on pages 467-469, DOI: 10.11646/zootaxa.4755.3.2, http://zenodo.org/record/3736817, {"references":["Corbet, G. B. & Hill, J. E. (1992) The Mammals of the Indomalayan Region: A systematic Review. Oxford University Press, Oxford, 488 pp.","Hill, J. E. (1965) Asiatic bats of the genera Kerivoula and Phoniscus (Vespertilionidae), with a note on Kerivoula aerosa Tomes. Mammalia, 29, 524 - 556. https: // doi. org / 10.1515 / mamm. 1965.29.4.524","Heaney, L. R., Dolar, M. L., Balete, D. S., Esselstyn, J. A., Rickart, E. A. & Sedlock, J. L. (2010) Synopsis of Philippine Mammals. Available from: http: // www. fieldmuseum. org / philippine _ mammals / (accessed 8 May 2019)","Heaney, L. R., Balete, D. S. & Rickart, E. A. (2016) The Mammals of Luzon Island: Biogeography and Natural History of a Philippine Fauna. Johns Hopkins University Press, Baltimore. xi + 287 pp.","Kingston, T., Jones, G., Akbar, Z. & Kunz, T. H. (1999) Echolocation signal design in Kerivoulinae and Murininae (Chiroptera: Vespertilionidae) from Malaysia. Journal of Zoology, 249, 359 - 374. https: // doi. org / 10.1017 / S 0952836999009917","Schmieder, D. A., Kingston, T., Hashim, R. & Siemers, B. M. (2010) Breaking the trade-off: rainforest bats maximize bandwidth and repetition rate of echolocation calls as they approach prey. Biology Letters, 6, 604 - 609. https: // doi. org / 10.1098 / rsbl. 2010.0114","Payne, J., Francis C. M. & Phillipps, K. 1985. A field guide to the mammals of Borneo. Sabah Society, Kota Kinabalu, 332 pp.","Kingston, T., Boo Liat, L. & Akbar, Z. (2006) Bats of Krau Wildlife Reserve. Penerbit Universiti Kebangsaan Malaysia, Bangi, 141 pp.","Rossiter, S. J., Zubaid, A., Mohd-Adnan, A., Struebig, M. J., Kunz, T. H., Gopal, S., Petit, E. J. & Kingston, T. (2012) Social organization and genetic structure: Insights from codistributed bat populations. Molecular Ecology, 21, 647 - 661. https: // doi. org / 10.1111 / j. 1365 - 294 X. 2011.05391. x","Nurul-Ain, E., Rosli, H. & Kingston, T. (2017) Resource availability and roosting ecology shape reproductive phenology of rain forest insectivorous bats. Biotropica, 49, 382 - 394. https: // doi. org / 10.1111 / btp. 12430","Schmieder, D. A., Kingston, T., Hashim, R. & Siemers, B. M. (2012) Sensory constraints on prey detection performance in an ensemble of vespertilionid understorey rain forest bats. Sensory constraints on prey detection performance in an ensemble of vespertilionid understorey rain forest bats. Functional Ecology, 26, 1043 - 1053. https: // doi. org / 10.1111 / j. 1365 - 2435.2012.02024. x"]}

Kerivoula whiteheadi Thomas, 1894 Type locality: Molino, Isabela Province, Luzon. Specimens examined. Philippine specimens examined are listed in Appendix A. Distribution. Endemic to the Philippines (Bohol, Camguin Norte, Cebu, Lubang, Luzon, Mindanao, Mindoro, Palawan, Sicogon, Siquijor and Tumaguin; Fig. 2). The type locality “Molino” probably refers to a currently nonexistent village on the east bank of the Ilagan River downstream from the city of San Mariano (M. van Weerd, pers. com., 6 May 2019). Description. Total length 67–86 mm, tail 29–41mm, ear 12–18 mm, forearm 29–33 mm, mass 2.5–5.5 g (Table 1). A clear north-south gradient in size exists from Camiguin Norte Island (north of Luzon, largest) to southern Mindanao (smallest; Tables 1 and 4). A small, delicate bat (Fig. 12A) with funnel-shaped ears, a long, slender tragus, and simple nostrils (Fig. 12C). The dorsal fur varies from dark brown (on Luzon) to reddish brown (on Mindanao), with the ventral fur somewhat paler (Fig. 12A, B). Each hair in the dorsal pelage has three bands (dark gray-brown base, light buff mid-band, and rufus-brown to bright rufus tips (Fig. 12D). The ventral hairs are also banded with a dark gray-brown base and light brown tips (Fig. 12E). The wing and tail membranes are dark. A scattering of a few short hairs is present along the posterior margin of the interfemoral membrane, but there is no definite fringe. Thick hair is present along the dorsal tail membrane, along the legs and on the feet (Fig. 12B). Skull as shown in Figure 13. GTL12.16–14.60, CCL 11.08–13.66 mm, GBB 6.42–7.14 mm, BH 5.15–6.09 mm, PC 2.92–3.26, C–M 3 4.89–5.94 mm, M 2 –M 2ext 4.31–5.11 mm, C 1 –C 1 1.36–1.93 mm, C–M 3 5.23–6.32 mm, MDL 9.22–11.24 mm (Table 1). The skull of K. whiteheadi (Fig. 13) is small and the braincase is highly inflated, producing a pronounced forehead. The rostrum narrows anteriorly, and the upper toothrows converge moderately. The center of the dorsal surface of the rostrum often has a ventral depression along the midline, above the level of the molars. The cochlea are large, and the basioccipital region is narrow. The height of the second upper incisor is half or slightly more than half of the first incisor. The first and second upper premolars, and all three lower premolars, are of nearly equal crown area, and are strongly laterally compressed (Fig. 7C). The lingual shelves of M 1 and M 2 are relatively small, leaving a broad embrasure at their posterior margins. The first lower incisor usually has three cusps, the second has three cusps, and the third usually has three cusps but occasionally two; they overlap substantially (Fig. 7D). Comparisons. Kerivoula whiteheadi differs from K. pellucida as noted above. Kerivoula whiteheadi and K. “ hardwickii A” are similar in size (average forearm length 31.1 and 33.0 mm, CCL 11.9 and 11.6 mm, respectively), but where they overlap geographically, K. hardwickii are slightly larger. K. whiteheadi is smaller overall than K. “ hardwickii B” (forearm length 35.0 mm, CCL 13.2 mm). The dorsal pelage of K. whiteheadi has a reddish tone (Fig. 12B, D), rather than medium or dark brown in the Philippine K. hardwickii. Both species have hairs that are dark at the base on the dorsum and venter with color bands—three dorsally and two ventrally (Fig. 12D, E). K. whiteheadi overall appears more bicolored than Philippine K. hardwickii with a rufus-brown dorsum and grayish-brown venter, whereas Philippine K. hardwickii appears brown overall (although there is some variation). A fringe of short, inconspicuous hairs along the posterior edge of the interfemoral membrane is present on Philippine K. hardwickii, but absent on K. whiteheadi (Fig. 12A). The wing attaches at a similar point below the outside toe of both species (Fig. 12A). The penis of male K. whiteheadi does not widen distally and lacks a characteristic nub that is present on the penis of males in Philippine K. hardwickii (J. Sedlock, pers. obs.). The skulls of K. whiteheadi usually have a more abrupt forehead than Philippine K. hardwickii. The latter often has a slight sagittal crest posterior to the interorbital region, a structure that is absent in K. whiteheadi (Figs. 8, 13). The depression along the midline of the rostrum in K. whiteheadi is absent in Philippine K. hardwickii. The teeth of Philippine K. hardwickii are proportionately substantially larger and more robust than those of K. whiteheadi (Fig. 10A, B). In Philippine K. hardwickii, the second upper incisor is about one-third the height of the second incisor; in K. whiteheadi, the second incisor is half or more the height of the first incisor (Figs. 7C, D, 10A, B). Finally, Philippine K. hardwickii lack the conspicuous lateral compression of the premolars present in K. whiteheadi. Kerivoula whiteheadi differs from K. papillosa in being much smaller overall in size (mean total length 79.9 mm and forearm length 31.1 mm, rather than 101.6 mm and 42.5 mm, respectively, mean CCL 11.99 mm rather than 15.78 mm; Table 1, 3; Heaney et al. 2010; unpublished data). The dorsal pelage of K. whiteheadi has a reddish tone with a distinct banding patterns on hairs (Fig. 12), rather than medium or dark brown with no clear banding pattern (but darker at base). K. whiteheadi ’s skull is small and delicate lacking a sagittal crest (Fig. 13), rather than larger and more robust with a sagittal crest (Fig. 9). Premolars are elongate (Fig. 7C, D), rather than round (Fig. 10C, D). Echolocation. The echolocation calls of K. whiteheadi in the Philippines are similar to those of K. pellucida in that they are broadband, high frequency, and short duration (Fig. 11; Table 5). Peak frequency of K. whiteheadi calls is 138 kHz. However, as stated above, K. whiteheadi calls are significantly shorter in duration and have a narrower bandwidth than those of K. pellucida (Table 5). There were no significant differences in call attributes among populations from Bohol, Cebu and Siquijor islands; however, we had sufficient recordings from only those three central Philippine islands and not from Luzon and Mindanao, which represent the largest and smallest individuals within the species, respectively. Ecology. Kerivoula whiteheadi exhibits a broad tolerance of vegetation types and elevations in the Philippines, from highly disturbed and fragmented lowland forest near sea level, to pristine montane forest at 1465 m above sea level (Sedlock 2002; Sedlock et al. 2008, 2011, 2014; Heaney et al. 2016). It has also been reported in karst areas within disturbed forest on limestone in the central Philippines and on Palawan Island (Esselstyn et al. 2004; Sedlock et al. 2014). Pregnant females have been captured in April, June and August, suggesting an asynchronous reproductive period as in Kerivoula species in Peninsular Malaysia (Nurul-Ain et al. 2017). No reports of roosting behavior exist from the Philippines. The authors have observed its slow and maneuverable flight, and its keen ability to quickly find and retreat into hollows or holes when released within a flight tent or room. For example, one individual flew up the leg of a pair of pants hanging over a line within minutes of release. When perching on the net wall of the flight tent, K. whiteheadi curls up with its head upright rather than hanging down (Sedlock 2002)., Published as part of Sedlock, Jodi L., Heaney, Lawrence R., Balete, Danilo S. & Ruedi, Manuel, 2020, Philippine bats of the genus Kerivoula (Chiroptera: Vespertilionidae): Overview and assessment of variation in K. pellucida and K. whiteheadi, pp. 454-490 in Zootaxa 4755 (3) on pages 473-478, DOI: 10.11646/zootaxa.4755.3.2, http://zenodo.org/record/3736817, {"references":["Thomas, O. (1894) Descriptions of two new bats of the genus Kerivoula. Annals and Magazine of Natural History, 14, 460 - 462. https: // doi. org / 10.1080 / 00222939408677834","Heaney, L. R., Dolar, M. L., Balete, D. S., Esselstyn, J. A., Rickart, E. A. & Sedlock, J. L. (2010) Synopsis of Philippine Mammals. Available from: http: // www. fieldmuseum. org / philippine _ mammals / (accessed 8 May 2019)","Sedlock, J. L. (2002) Autecology and the conservation of insectivorous bats on Mt. Makiling, Philippines. Silliman Journal, 42 (1), 163 - 201.","Sedlock, J. L., Weyandt, S. E., Cororan, L., Damerow, M., Hwa, S. & Pauli, B. (2008) Bat diversity in tropical forest and agropastoral habitats within a protected area in the Philippines. Acta Chiropterologica, 10 (2), 349 - 358. https: // doi. org / 10.3161 / 150811008 X 414926","Sedlock, J. L., Ingle, N. R. & Balete, D. S. (2011) Enhanced sampling of bat assemblages: a field test on Mount Banahaw, Luzon. Fieldiana: Life and Earth Sciences, 2, 96 - 102. https: // doi. org / 10.3158 / 2158 - 5520 - 2.1.96","Heaney, L. R., Balete, D. S. & Rickart, E. A. (2016) The Mammals of Luzon Island: Biogeography and Natural History of a Philippine Fauna. Johns Hopkins University Press, Baltimore. xi + 287 pp.","Esselstyn, J. A., Widmann, P. & Heaney, L. R. (2004) The mammals of Palawan Island, Philippines. Proceedings of the Biological Society of Washington, 117, 271 - 302.","Nurul-Ain, E., Rosli, H. & Kingston, T. (2017) Resource availability and roosting ecology shape reproductive phenology of rain forest insectivorous bats. Biotropica, 49, 382 - 394. https: // doi. org / 10.1111 / btp. 12430"]}

Heaney, Lawrence R., Balete, Danilo S., Rickart, Eric A., Veluz, Maria Josefa, Jansa, Sharon A. (2014): Three New Species of Musseromys (Muridae, Rodentia), the Endemic Philippine Tree Mouse from Luzon Island. American Museum Novitates 2014 (3802): 1, DOI: 10.1206/3802.1, URL: http://www.bioone.org/doi/abs/10.1206/3802.1

Balete, Danilo S., Rickart, Eric A., Heaney, Lawrence R., Alviola, Phillip A., Duya, Melizar V., Duya, Mariano Roy M., Sosa, Timothy, Jansa, Sharon A. (2012): Archboldomys (Muridae: Murinae) Reconsidered: A New Genus and Three New Species of Shrew Mice from Luzon Island, Philippines. American Museum Novitates 2012 (3754): 1-60, DOI: 10.1206/3754.2, URL: http://www.bioone.org/doi/abs/10.1206/3754.2

Rafflesia verrucosa Balete, Pelser, Nickrent & Barcelona, sp. nov. (Figs. 2A–F, 3A–F) Rafflesiae baleteae floribus bisexualis, et R. leonardae antheris numerosis, 20-21, etiam R. aurantiae formis ramentorum similis. Autem verrucis prominentibus in perigonio et diaphragmate orae aperturae attingentes, disco cupulato processibus laminaribus sinuosis consociatis confertim impletis, corona pubiscentia densa ad trientum basilarem disci externi extensa, annulo latissimo toroido relative laevi, antheris minimus numerosioris ab omnibus aliis rafflesiis parvis praesertim differt. Type:— PHILIPPINES: Mindanao Island: Davao Oriental Province: Manay (also known as Man-ay) Municipality: Mt. Kampalili, 7.29112° N, 126.31520° E, 1470 m, 3 March 2010, Balete 17 (holotype PNH, isotypes CAHUP, F, L, SING, US). Endophytic holoparasite. Mature buds to 7.5 cm in diameter. Cupule 2.0 cm high to 6.0 cm wide. Bracts (or bud scales) numerous in three to four imbricate whorls, outermost smallest, ca. 1.6 cm long, 1.5 cm wide, innermost largest to ca. 5.1 cm long, 6.5 cm wide. Flowers 14.5–16 cm in diameter when fully expanded, 11.5–13 cm high (Fig. 2A), with a very mild putrescent smell when fresh. Perigone tube ca. 3.5–5.5 cm long, inner surface with minute pustules (broccoli-like), with a midlayer of sclerenchymatous tissue that extends to the perigone lobes, this persisting beyond senescence and into fruit maturity, often forming a cup-like structure on top of the maturing fruit (Fig. 2B). Perigone lobes 5, orbicular to broadly orbicular, 4.2–4.5 cm long, 6.5–8.0 cm wide, margins irregularly sinuate, reddish orange or cinnamon, becoming dark brown with age; adaxial surface verrucose, warts prominently raised, solitary, irregular in shape, size, and density, usually roundish, less often rod-like to narrowly elongated, white-tipped in newly opened flowers, with age becoming concolorous with perigone adaxial surface, abscising upon flower senescence leaving a persistent wood-like tissue extending from the perigone tube (Fig. 2B); abaxial surface covered with hardened tissue that is irregularly cracked and partially flakes off (sometimes already early in the development of the flower) revealing a pitted inner layer (Fig. 2D). Diaphragm 7–9 cm in diameter, 2–3.5 cm wide from aperture rim to base of perigone lobes, 9–11 mm thick at base, becoming thinner towards the aperture; concolorous with or slightly darker than the adaxial perigone surface; densely covered with prominently raised, pleated, plate-like warts that are white-tipped in newly opened flowers, these variable in size, larger ones 3–8 mm tall, intermixed with smaller ones, ca. 1–2 mm tall (Fig. 2C); windows absent; aperture 3.5–4 cm in diameter. Disk 5–5.5 cm in diameter, ca. 1–1.2 cm thick midway between the margin and the point where the disk joins the column, cup-shaped with a prominently raised margin, concolorous with diaphragm; processes forming an interconnected system of tightly packed, laminar plates with erose margins, ca. 3–11 mm tall (Fig. 2C, E–F); column very short, up to ca. 1 cm from the floor of the floral tube to the upper surface of disk, ca. 2.3 cm wide, deeply grooved, number of sulci corresponds to the number of anthers, septa between sulci ca. 1 cm tall, with acicular hairs (‘bristles’ sensu Nais 2001) ca. 1.5 mm long; annulus ca. 7–10 mm wide, ca. 5–6 cm in diameter, doughnut-shaped (toroid), concolorous with the rest of the perigone tube, surface shallowly pitted and rugose (Fig. 2F). Ramenta to 7 mm long, covered with clavate pustules, polymorphic, filiform to variously branched or cleaved apically, those at the floor of the perigone tube longer and denser becoming shorter and more widely spaced on the diaphragm, nearly absent near the aperture rim, white-tipped in newly opened flowers, becoming concolorous with perigone abaxial surface with age. Flowers bisexual (Fig. 3A); anthers 20 (or 21), globular, ca. 2 mm in diameter, prominently protruding from very shallow anther sulci that are 2–3 mm long, 3–4 mm wide (Fig. 3B), basal third of disk formed into a pubescent corona with bristle-like, acicular hairs to 0.5 mm (Fig. 2E, 3B); ovary ca. 1.5 cm tall, 4.5 cm wide, lenticular or reniform, becoming broadly and irregularly so as the ovary matures (Fig. 3C); developing fruits to 7 cm in diameter and 4.7 cm high, sclerenchymatous perigone tissue persistent. Distribution and habitat:— Rafflesia verrucosa is only known from the southeastern slope of Mt. Kampalili, Davao Oriental Province, Mindanao Island (Fig. 1). It is found between 1350 and 1550 m. Despite extensive search efforts in the area, it was not encountered at lower (900–1350 m) or higher (1550–1700 m) elevation, although the presence of Tetrastigma vines, which is the host for this species, was confirmed throughout the elevational range explored. The habitat of R. verrucosa is montane forest in relatively rocky and moderately sloping terrain. Leaf litter is ca. 2–5 cm thick. The canopy is ca. 15 m tall and emergents reach 20 m. The larger trees in this forest commonly reach a DBH up to ca. 90 cm. Dominant trees are oaks (Lithocarpus Blume 1826: 526), laurels (Litsea Lamarck 1792: 574), and myrtles (Syzygium Gaertner 1788: 166). Mosses are common on tree trunks and branches, and often also at the bases of trunks. Other epiphytes include ferns and lycophytes, Medinilla Gaudichaud-Beaupré (1830: 484), and orange-flowered Rhododendron Linnaeus (1753a: 392). Understory vegetation is abundant and composed of tree ferns, other ferns and lycophytes, ground orchids, gingers and grasses. Canopy vines such as Freycinetia Gaudichaud-Beaupré (1824: 509), climbing bamboos, rattans, Smilax Linnaeus (1753b: 1028), Tetrastigma, and other lianas are likewise common. Several R. verrucosa plants were found on Tetrastigma vine roots growing exposed on rocky ground which resulted in deformed buds and flowers. Etymology:— The specific epithet of this new taxon is derived from the Latin verruca (wart), which calls attention to the unique, prominently raised warts on the perigone lobes and diaphragm. Ecology:— Rafflesia verrucosa is restricted to plants of a single, presently unidentified species of Tetrastigma (Fig. 3D, specimen not collected). Flowers and buds were only observed on the roots of their host plants and are absent on prostrate and aerial stems. Tetrastigma roots bearing Rafflesia flowers or scars range from 6 to at least 25.5 mm in diameter and buds were found on roots buried up to 7 cm below ground level. A total of 15 clusters of flowers and buds were discovered in an area of ca. 20 m × 1 km. At one site, at least 30 Rafflesia buds in different stages of development as well as senescent flowers were observed (Fig. 3E). This is equivalent to ca. 7–8 flower clusters per hectare. At the time of discovery of R. verrucosa (February–March, 2010), a notably large number of senescent flowers and developing fruits were observed. This suggests that flowering must have peaked around October to December similar to what is reported for R. mira, a species found in the adjacent Mt. Candalaga, Compostela Valley (Madulid et al. 2006). Since some mature buds, newly opened flowers, and early senescent flowers were also observed during our visit in February–March, flowering appears to continue intermittently throughout the year. Several buds as well as a maturing ovary of a senescent flower showed evidence of damage by animals, mainly as bite marks of unidentified small-toothed mammals (Fig. 3F). Systematic trapping in the area where R. verrucosa occurs yielded at least ten species of native non-volant small mammals, including a gymnure (Podogymnura), a shrew (Crocidura beatus), a tree shrew (Urogale everetti), two shrew-mice (Crunomys), a moss-mouse (Tarsomys), an arboreal tree-mouse (Apomys), a forest mouse (Apomys), the large Mindanao forest rat (Bullimus bagobus) and the common Philippine forest rat (Rattus everetti). Also reported in the area by local hunters are two species of squirrels (Exilisciurus concinnus and Sundasciurus philippinensis). Likewise, we also documented the presence of larger mammals, including palm civet (Paradoxurus hermaphroditus), Philippine warty pig (Sus philippinensis) and Philippine brown deer (Cervus mariannus). At ca. 1500 m, wild pig diggings on the ground around three populations of R. verrucosa are extensive but no sign of feeding on the buds and flowers was observed, although several were trampled upon. The role of these mammals in Rafflesia dispersal in the Philippines remains unstudied. But in Borneo, for instance, both squirrels (Callosciurus) and tree shrew (Tupaia) are known to feed on Rafflesia fruits (Emmons et al. 1991). Conservation:— Rafflesia verrucosa plants grow in the tropical montane forest formation (Fernando et al. 2008) which is increasingly undergoing disturbance and fragmentation from clearing for abaca (Musa textilis Née 1801: 123) plantations by native Mandayas. In addition, trees in the area are cut to harvest the young palm heart of the rattan plants that climb these trees. This practice resulted in areas of up to 100 m 2 in which tall trees have been cleared and the understory has been damaged. Despite this disturbance, the forest of the Mt. Kampalili Range is relatively intact and certainly does not suffer from the same degradation and loss that plague the lowland forest in which the majority of the Philippine Rafflesia are found (Barcelona et al. 2009b). Although R. verrucosa is moderately common in the area, it is not known from any other site on Mt. Kampalili or elsewhere and we therefore recommend that the clearing of the forest for abaca plantations and the harvesting of other forest products must be carefully managed in the wider area to ensure the continued survival of this new species of Rafflesia. We strongly support efforts by local people (e.g. Taocanga Tribal Council Association), environmental organizations (e.g. Philippine Eagle Foundation) and concerned government agencies (e.g. Department of Environment and Natural Resources and the Department of Tourism Region XI) to place certain portions of Mt. Kampalili under protected area and ancestral domain management. Additional specimens examined (paratypes):— PHILIPPINES: Mindanao Island: Davao Oriental Prov., Manay (= Man-ay) Municipality: Mt. Kampalili, 7.29796°N, 126.31216°E, ca. 1378 m, 25 February 2010, Balete 16 (SING); 7.29537°N; 126.31606°E, ca. 1550 m, 1 March 2010, Balete 18 (PUH, US).