Description of a New Large-Bodied Species of Apomys Mearns, 1905 (Mammalia: Rodentia: Muridae) from Mindoro Island, Philippines
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 108(2):302-318. 1995. Description of a new large-bodied species of Apomys Mearns, 1905 (Mammalia: Rodentia: Muridae) from Mindoro Island, Philippines Luis A. Ruedas Department of Biology, University of Puerto Rico, Cayey University College, Avenida Antonio R .. Barcelo, Cayey, Puerto Rico 00736 Abstract. -Apomys gracilirostris, a new species in the Philippine endemic murid. genus Apomys, is described from the Mount Halcon highlands on Min doro Island. The new species is compared with its larger-bodied congeners: A. abrae, A. datae, A. sacobianus, and the two nominal subspecies of A. insignis, A. i. insignis and A. i. bardus. Twenty-eight cranial, mandibular, and dental measurements were taken on the 83 specimens representing these taxa and the new species. All species examined formed discrete groups in a principal com ponent analysis; there were no apparent differences between the two nominal subspecies of A. insignis. The new species differs from congeners in its com bination of large body, very long tail, dark pelage, gracile cranium with a long rostrum, and highly compressed upper incisors; the most diagnostic features of the species are the extremely fine upper and lower incisors (with no enamel on the lower), and the degree of molar reduction and simplification. In 1905, Edgar A. Mearns proposed the et al. 1993), A. microdon from southern Lu generic appellation Apomys for represen zon and Catanduanes Islands, A. musculus tatives of a new species of rodent he had from Luzon and Mindoro Islands, and A. collected on Mount Apo, in southern Min datae and A. abrae from Luzon Island. An danao Island, Philippines (Mearns 1905). additional species, currently undescribed, Different investigators historically have recently has been reported from a number shown considerable flux with respect to the of islands iil""--the central Visayas group of taxonomic placement of the genus (review the Philippine Archipelago (Musser 1982, by Musser 1982). However, since the ex Musser & Heaney 1992; L. R. Heaney, pers. haustive review and diagnosis of Apomys comm.). by Musser ( 1982), there has been no doubt During a biodiversity inventory survey as to the validity of these organisms con carried out in 1992 on some components of stituting a genus level category. As currently the major Pleistocene island masses (sensu defined (Musser 1982, Musser & Heaney Heaney -1986), one of the localities that was 1992), Apomys is restricted to the Philip visited was the Mount Halcon massiff, in pine Archipelago (excluding the Sunda Shelf the northern part of the island of Mindoro. island of Palawan and associated peripheral Mount Halcon, reported to reach ca: 2500 islands). m in elev.ation, had been explored by Edgar Eight species are now included in the ge A. Mearns in 1906 (Merrill 1907a); how nus: A. hylocoetes and A. insignis from Min ever, only 17 of the 496 ma·mmals brought danao Island, A. littoralis from Mindanao, back by him were from Mindoro. Most of Leyte, Biliran, and Bohol Islands (Heaney his collection from the Mt. Halcon area con et al. 1989, Musser & Heaney 1992, Rickart sisted of Crocidura; all told, eight nominal VOLUME 108, NUMBER 2 303 taxa in four orders were collected by him, case (BB), breadth across incisive foramina including the holotypes of five nominal taxa. (BIF), breadth across incisor tips (BIT), The object of the current exploration of the breadth of palatal bridge at first, and at third Mount Halcon area was to increase sample molars (BMl, BM3), breadth ofmesoptery sizes of some of these taxa in order to arrive goid fossa (BMF), breadth of rostrum (BR), at a better assessment of geographic and breadth of zygomatic plate (BZP), height, nongeographic variation, as well as to de and length, of auditory bulla (HB, LB), termine whether additional species, which height of braincase (HBC), interorbital Mearns may have missed, also existed in breadth (IB), incisive foramina to M 1 the area. (IFM 1; modified to be straightline shortest One mammal species found between 12 5 5 distance, rather than anteroposterior dis m and 1900 m elevation on the north slope tance), length of diastema (LD), length of of Mount Halcon was a medium-sized ro incisive foramina (LIF), alveolar length of dent, similar to Apomys in having a slender maxillary toothrow (LMT), length of nasals rostrum, as opposed to the short, high, blunt (LN), length of palatal bridge (LPB), length rostrum of Rattus. The lower incisors were of rostrum (LR), occipitonasal length (ONL), more procumbent than those of Rattus, and palatal length (PL), postpalatal length (PPL), lacked pigment. The general gestalt, mostly and zygomatic breadth (ZB). Five addition related to how gracile this rodent was, there al measurements not ·Used by Musser (1979) fore pointed, barring the relatively large size, were taken: length of dentary (LMAND)~ to its pertenence in the genus Apomys. Sub the greatest length of dentary from upper sequent examination of cranial morpholo incisive alveolar rim to mandibular con gy, and conversations with colleagues, con dyle; length of mandibular toothrow (LMT), firmed this suspicion. Accordingly, individ alveolar length of mandibular toothrow; uals collected in. that taxon are described height of mandible (HMC), greatest height below as a new species of Apomys from the of mandible; height of mandibular ramus northern highlands of-Mindoro. (HMR), lowest height of ramus in mandib ular incisive diastema; and thickness of mandible (TM), measured at thickest point Methods below m2. Except for specimens of the new species Statistical analyses were performed on an (at the Cincinnati Museum of Natural His 80486 equipped 33 MHz microcomputer tory), and those of Apomys sacobianus (Na using the Statistical Analysis System soft tional Museum of Natural History, USNM), ware, version 6.03 (SAS Institute 1988a, all specimens examined are from the Field 1988b), generally following Ruedas & Bick Museum of Natural History (FMNH). Ter ham (1992). Univariate statistics, including minolo.gy referent to molar cusps follows mean, standard error, moment statistics Miller ( 1912) for upper molars, and Musser (skewness, g1; and kurtosis, g2) and tests of and Heaney (1992) for lower molars. Cra normality were performed invoking the nial and mandibular measurements were ''normal'' option of the UNIVARIA TE taken with a digital caliper to the nearest procedure, which tests for normality using 0.01 mm. Twenty-eight cranial, mandibu the Shapiro-Wilk statistic, W, and provides lar, and dental measurements, listed below, the associated probability value. A principal were taken; the 23 cranial measurements component analysis was carried out to de were described and illustrated in Musser termine whether any group separation oc (1979); mandibular measurements are de curred using the measurements taken. Such fined below. Measurements (abbreviations a posteriori grouping methods are preferred in parentheses) included: breadth of brain- herein over a priori grouping methods (mul- 304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. - Values for the first through third eigen A. datae: Philippines.: Luzon Island, vectors from the principal component analysis carried Mountain Prov.; Mount Data, 5300-8000 out on the correlation matrix of the cranial and man ft. FMNH 62695, 62696, 62699, 62720, dibular morphometric data of specimens of Apomys. Refer to text for character abbreviations. The first four 62741, males; 62706, 62709, 62711, 62712, principal components together account for 91.5°/o of 62725,62727,62731,62733-62735,62742, the variation (68.5°/o, 13.3°/o, 5.5°/o, and 2:2°/o, respec 62744, females. Mountain Prov.; no specific tively). Results.ofthe principal component analysis are locality. FMNH 62761, female. graphically summarized in Fig. 1. A. insign.is bardus: Philippines: Mindanao Component Isl., Zamboanga del Norte Prov.; Grand 1 2 3 Malindang Mountains·, Dapitan Peak, 6700- 7450 ft. FMNH 87568 (young), 87571- BB 0.2092 - 0.2372 -0.0489 87573, males, 87567, 87569, 87570, fe BIF 0.1498 0.3402 0.0601 BIT 0.1244 0.2985 0.4763 males .. BMl 0.1684 - 0.0566 -0.5707 A. i. insignis: Philippines: Mindanao Isl., BM3 0.1942 - 0.2232 - 0.1403 Bukidnon Prov.; Malaybalay, Mount Ki BMF 0.0993 0.3870 - 0.0400 tanglad, 4200-5000 ft. FMNH 92806, BR 0.2279 -0.0040 0.2225 92808-92812,92815,92816,92818,92819, BZP 0.2509 0.1233 0.0151 HB 0.1408 0.3926 0.0275 92821,males; 92807, 92813, 92814, 92817, HBC 0.2454 - 0.0082 - 0.0001 92820, 92822, females. Davao Prov.; East IB 0.0362 -0.1522 0.3550 slope of Mount McKinley, 3500-7100 ft. IFMl 0.1443 - 0.3582 0.0222 FMNH 56267, 56270, 56276-56278, males; LB 0.2012 0.0836 - 0.2576 56266,56268,56269,56272-56275, 56282, LD 0.2465 -0.1 715 - 0.0298 LIF 0.2353 0.1279 -0.2145 56285, 56288, females. LM13 0.2081 0.2802 -0.1150 A. sacobianus: Philippines: Luzon Isl., LN 0.2610 - 0.0246 0.0379 Pampanga Prov.; Angeles, Clark Air Base, LPB 0.2370 -0.1172 0.1331 Sacobia River. USNM 304352, 557717. LR 0.2453 -0.1326 0.1170 Apomys n. sp.: see below, holotype and ONL 0.2653 -0.0583 0.0164 PL 0.2651 0.0501 - 0.0369 paratypes. PPL 0.2576 - 0.0188 0.0451 ZB 0.1841 -0.2135 0.2830 Results Descrip ti ve statistics for species of Apo- - - mys examined in this study are listed in Appendix 1. Results of the principal com tiple range tests, canonical discriminant ponent analysis (Fig. 1) show considerable, analysis) because there is no prior hypoth non-overlapping variation among different esis as to the putative identity of the spec nominal taxa within Apomys. The first four imens being examined. principal components account for 91.5°/o of the variation (Table 1). Principal compo Specimens Examined nent 1 accounts for · 68°/o of the variation A.