Tabu 1. Numbers of nonvolant small mammals captured in traps in heavily disturbed lowland agricultural area (Site 3) secondary lowland forest (Site 4). disturbed lower montane forest (Site 5), primary mossy forest (Site 6) primary montane forest (Site 7), and lower mossy forest (Site 8) on Camiguin Island. The numbers of captures per KM) trap-nights are given in parentheses. See Heaney and Tabaranza (2005a) for full site descriptions. Asterisks mark species presumed to be present; see Methods. common in disturbed lowland forest at 1000 m (Site 4) and uncommon in disturbed lower montane forest at 1000-1300 m and mossy forest between 1200 and 1400 m elevation (Sites 5 and 6). Limited netting did not detect this species in montane primary forest at 1275 m elevation (Site 7). The occurrence of C. brachyotis in these habitats is consistent with records from other Philippine islands; for example, on Catanduanes, Leyte, Luzon, and Negros, this species was most abundant in agricultural land and secondary forest (Heaney et al, 1989, 1991, 1999; Heideman & Heaney, 1989; Ingle, 1992; Rickart et al, 1993), similar to Sites 1 and 3 to 5 on Camiguin. Records from the 1960s indicate that the species occurs along the entire elevational gradient, from sea level to near the peaks, although most specimens are from below 1000 m (ie, below the transition to montane forest; Fig. 2). Eight adult females (mean 295 472 g) taken in May 1992 and 1994 were pregnant with single embryos (CRL 5-26 mm). Two of the pregnant females (22 and 35 g) and three non-pregnant ones (325-39 g) were lactating. Adult females with enlarged mammae but not pregnant or lactating had mean weight of 292 23 g (range 26 332 g, N 13). Five nulliparous females had a mean weight of 165 g 20 g (range 14-19 g). Males with abdominal testes had a mean weight of 274 26 g (range 22325 g, N 15). On Negros Island, this species probably produces two young per year, one in March-april and another in August-september (Heideman, 1995). Females are slightly larger than males in most external and cranial dimensions (Table 4), a trend quite similar to specimens from Mt. Kitanglad, Mindanao (Heaney et al, unpubl. data). Specimens from Biliran, Leyte and Maripipi, in contrast, showed the opposite trend (Rickart et al, 1993). Specimens Examined — Total 91. Site 1 (27 fmnh); Site 3 (5 ms-iit); Site 4 (4 fmnh, 14 msu-iit); Site 6 (1 fmnh, 3 su-iit); Site 1112 dmnh); Site 12 (3 dmnh); Site 13 (6 dmnh); Site 16 (2 rom); Site 17 (14 rom). Harpyionycteris whiteheadi Thomas, 1 896 The harpy fruit bat has been reported previously from Camiguin (Peterson & Fenton, 1970), as well as from Marinduque, Masbate, Mindoro, Negros, southern Luzon, and throughout Greater Mindanao (Heaney et al, 1998, 1999). It is generally restricted to primary or lightly disturbed forest; it is usually rare in lowland forest but often is moderately common in montane forest from roughly 800 m to at least 1800 m. It apparently feeds heavily on the fruits of vines pandans (Freycinetia sp) and figs (Ficus spp); (Heaney et al, 1989, 1999; Heideman & Heaney, 1989; Rickart et al, 1993). In May 1992, we netted one pregnant female weighing 123 g with a single embryo (CRL 28 mm) at 10 m elevation, in a lowland 32 FIELDIANA: ZOOLOGY


e a Ctf 00 60 s 5 SI es C-o T3 a ; oo He o o oo on j P t3 a " c O oo 2 c5 S E u Q BO S c 6 5SE S of Q2 a i g — 6 E c a B Z 1 E HIS 00 — r. t-. 4" "3" IT) IT) Tfr OS OS CTS -4SO rn j A, — I CM in O s— r o 11 in in SO O (N — N • ; N — I — N so so oo' so O in > in I I 0O VO -Tt n n r r I in SO CO N CO Sa3 X in CO in o — OS 1 O OO o s — 1 so 4 Sh S u. -5 5 sj agricultural area (Site 1; Table 3). On Mindanao (Kitanglad Range), pregnant females were similarly recorded in May, with further records of pregnancy in April, August, and October; lactation was noted in March, April, and August (Heaney et al, unpubl. data). In March 1995, at Site 7 (1275 m in primary montane forest), where Freycinetia sp. were abundant, we often heard the distinctive Harpyionycteris whistles (Rickart et al, 1993; Heaney et al, 1999), but none were captured. Combined with records from the 1960s (Fig. 2), these data indicate that it occurs on this island from sea level to 1500 m in lowland, montane, and mossy forest. External and cranial measurements of H. whitehead on Camiguin are comparable to those from Mindanao (Mt. Kitanglad) but are slightly larger than those found on Leyte and Luzon (Mt. Isarog) (Heaney, 1984; Rickart et al, 1993; Heaney et al, 1999, unpubl. data). Specimens Examined — Total 4. Site 1 (1 fmnh); Site 11 (1 dmnh); Site 12 (1 dmnh); Site 16 (1 ROM). Macroglossus minimus (E. Geoffroy, 1810) The dagger-toothed flower bat occurs from Thailand to Australia and is found throughout the Philippines (Heaney et al, 1998). Within the Philippines, it occurs in virtually every habitat in the country, from sea level to at least 2250 m. It is often abundant in agricultural and heavily disturbed areas, is common in secondary forest, and usually is uncommon in primary forest. It is most often associated with domestic or wild banana (Musa spp; Heaney et al, 1989, 1999; Heideman & Heaney, 1989; Rickart et al, 1993). On Camiguin, our limited netting showed M. minimus to be abundant in a highly disturbed lowland agricultural area at ca. 10 m elevation (Site 1), present in a heavily disturbed lowland agricultural area at 100 m (Site 3), uncommon in disturbed lowland forest at 1000 m (Site 4), common in disturbed lower montane forest at 1000-1300 m (Site 5), and present in mossy forest between 1200 and 1400 m elevation (Site 6; Table 3). Limited netting did not detect this species in primary montane forest at 1275 m elevation (Site 7). Combined with prior records, it is apparent that this species occurs on Camiguin from sea level to at least 1200 m, in lowland and montane forest, in both disturbed and undisturbed forest (Fig. 2). HEANEY ET AL: THE MAMMALS OF CAMIGUIN ISLAND 33


in cranial and external measurements is slight, Camiguin specimens tend to cluster consistently within the upper ranges of most features measured (Table 4; Heaney and Rabor, 1982; Heaney et al, 1991, 1999, unpubl. data; Rickart et al, 1993). Specimens Examined — Total 56. Site 1 (15 fmnh); Site 3 ( 1 ms-iit); Site 4 (4 su-iit); Site 5 (8 su-iit); Site 6 (1 su-iit); Site 1112 dmnh); Site 13 (5 dmnii); Site 14 (2 dmnh); Site 17 (8 dmnii). Ptenochirus jagori ( Peters, 1861) The musky fruit bat is a common Philippine endemic, occurring throughout the archipelago with the exception of the Batanesbabuyan and Palawan faunal regions from sea level to at least 1800 m (Heaney et al, 1998). Our limited netting on Camiguin showed Ptenochirus jagori to be common in a lowland agricultural area at 10 m elevation (Site 1), less common in disturbed lowland forest at 1000 m (Site 4), and scarce in primary mossy forest at 1300 m (Site 6; Table 3). Combined with records from the 1960s, these data indicate that the species is widespread in lowland and montane rain forest from sea level to at least 1200 m (Fig. 2), though probably its abundance declines with increasing elevation and with increasing levels of disturbance (Table 3), as noted elsewhere (Heaney et al, 1989, 1999, unpubl. data; Heideman & Heaney, 1989; Rickart et al, 1993; Lepiten, 1997). Three adult females, netted in May 1992 and 1994, weighing an average of 70 g (range 6875 g), were pregnant with a single embryo each (CRL 5 10 mm). Three nonpregnant females with large mammae had an average weight of 735 g (range 72-74 g), and two nulliparous females weighed 67 and 685 g. Eleven adult males had a mean weight of 732 47 g (range 64-78 g, N 11). Pregnant females of P. jagori have been recorded also in May on Luzon (Mt. Isarog) (Heaney et al, 1999). On Mindanao (Kitanglad Range), pregnant females were recorded in March, May, July, and August and lactating females in May to June and August to December (Heaney et al. unpubl. data). Heideman and Powell (1998) found that on Negros Island, P. jagori gives birth to a single young twice each year: the first in late March or early April and the second in August. It was further discovered that this species undergoes delayed implantation and early development that lasts for five months, shorter than in two other endemic species of Philippine pygmy fruit bats, Haplonycteris fischeri and Otopteropus cartilagonodus, where the phenomenon was first detected (Heideman, 1989; Heideman et al, 1993; Heideman & Powell, 1998). Additionally, this condition was apparently exhibited by primiparous young adult females only, allowing them to give birth only once in their first year, which had the effect of enabling them to synchronize breeding with the adult females the following year (Heideman & Powell, 1998). Males are somewhat larger than females in most cranial and external dimensions, as on Biliran, Leyte, and Maripipi (Table 4; Rickart et al, 1993). Cranial and external measurements (Table 4) are noticeably larger than those for series from Catanduanes and southern Luzon and were similar to those from Biliran, Leyte, and Maripipi (Heaney, 1984; Heaney et al, 1991, 1999; Rickart et al, 1993). Specimens Examined — Total 46. Site 1 (8 fmnh); Site 416 ms-iit); Site 6 (1 su-iit); Site 11 (6 dmnh); Site 13 (6 dmnh); Site 17 (9 rom). Pteropus hypomelanus Mearns, 1905 The common island flying fox occurs from Thailand to Australia and is found throughout the Philippines with the exception of the Palawan faunal region. It is often common in agricultural areas from sea level to ca. 900 m and is absent in primary forest (Heideman & Heaney, 1989; Heaney et al, 1991, 1998; Utzurrum, 1992; Rickart et al, 1993). Records from the 1960s document it from Camiguin at elevations from about 250 to 1500 m (Fig. 2), but we netted none in the 1990s; because this species typically flies above the canopy and our nets were set not more than about 4 m above the ground, our failure to catch any does not necessarily indicate any change in their abundance. External and cranial measurements show only slight variations with those of specimens from Dinagat, Panay, Leyte, and Maripipi (Heaney & Rabor, 1982; Rickart et al, 1993). Specimens Examined — Total 8. Site 10 (2 dmnh); Site 12 (1 dmnh); Site 13 (2 dmnh); Site 17 (3 ROM). Pteropus pumilus Miller, 1910 The little golden-mantled flying fox is endemic to the Philippines (aside from a single population on Miangas Island. Indonesia, adjacent to Mindanao) and occurs throughout the archipelago, with the exception of the Batanesbabuyan 36 FIELDIANA: ZOOLOGY


and Palawan regions (Heaney et al, 1998). Previously reported from Camiguin as P. tablasi, the species was revised to include P. tablas and P. balutus as synonyms under Pteropus pumilus (Klingener & Creighton, 1984). It is associated with primary and good secondary lowland forest from sea level to about 1100 m, and it is uncommon outside of forest. Additionally, it is most common on small islands and is uncommon to rare on larger islands. Pteropus pumilus often is netted in clearings or on ridgetops (Heideman & Heaney, 1989; Utzurrum, 1992; Rickart et al, 1993). We did not encounter P. pumilus during the 1990s, but the 1967-1969 surveys obtained 42 individuals at four sites from ca. 250 to nearly 1000 m elevation (Fig. 2). As with Pteropus hypomelanus, this species usually flies above the canopy, so our failure to catch any in the 1990s does not necessarily indicate a change in their status on the island. Specimens Examined — Total 42. Site 11 (19 dmnh); Site 13 (2 dmnh); Site 14 (1 dmnh); Site 17 (20 ROM). Family Emballonuridae — Sheath-tailed Bats Emballonura alecto (Eydoux and Gervais, 1836) The Philippine sheath-tailed bat is a common cave-dwelling species that occurs throughout most of the Philippines and is also known from Borneo, the Moluccas, and Sulawesi (Koopman, 1989). It has been recorded only in lowland areas (450 m and below) in disturbed forest and agricultural areas with scattered remnant forest, with most captures recorded from caves, under large boulders, or in man-made tunnels (Heaney et al, 1991, 1999; Rickart et al, 1993). We did not record this species during our fieldwork in 1994 and 1995, but in 1967 specimens were taken from Tag-ibo Cave at 400 ft (ca. 120 m) and at 1400-3300 ft (ca. 400-1000 m) on Mt. Mambajao (see also Heaney, 1984). Comparison of external and cranial measurements with series from Leyte and Biliran shows little variation (Table 4; Rickart et al, 1993). Specimens Examined — Total 4. Site 17 (2 rom); Site 18 (2 rom). Family Rhinolophidae — Horseshoe-nosed Bats Rhinolophus arcuatus Peters, 1871 The arcuate horseshoe bat is widespread from Sumatra to New Guinea and throughout the Philippines (Heaney et al, 1998), including the Palawan faunal region (Esselstyn et al, 2004). It is most often encountered from lowlands to at least 1350 m in agricultural lands to primary lowland and montane forest and occasionally roosts in caves (Heaney et al, 1991, 1999; Rickart et al, 1993). Two groups that differ in size and habitat are recognized within this species: a smaller morphotype, designated R. arcuatus — s, that occurs in the lowlands or disturbed habitats, and a larger one, designated R. arcuatus — 1, found in forest at higher elevations (Ingle & Heaney, 1992). In May 1994, we netted this species at 1000 m elevation in disturbed lowland forest (Site 4). Of two adult females netted, one (15 g) was pregnant with one large embryo (CRL 28 mm), and the other was lactating (125 g). Pregnancies in this species were recorded also in March on Luzon (Mt. Isarog) and in April on Biliran, Leyte, and Maripipi (Rickart et al, 1993; Heaney et al, 1999). Cranial and external measurements (Table 5) of the Camiguin specimens are consistently larger than those in series from Biliran, Leyte, Luzon (Mt. Makiling), Maripipi, and Mindanao (Mt. Kitanglad), which all fall within the dimensions of the smaller morphotype, R. arcuatus — s (Ingle & Heaney, 1992; Rickart et al, 1993; Heaney et al, 1999, unpubl. data). Instead, their external and cranial dimensions are comparable to the larger morphotype, R. arcuatus — 1 (Ingle & Heaney, 1992). Systematics in this "species" are badly in need of detailed study. Specimens Examined — Total 10. Site 4 (4 msu-iit); Site 13 (6 dmnh). Rhinolophus inops K. Anderson, 1905 The Philippine forest horseshoe bat is common to abundant in primary lowland and montane forest from sea level to 2250 m and is usually rare in secondary forest and mossy forest (Heaney et al, 1991, 1998, 1999; Rickart et al, 1993). Improvements in our understanding of Philippine Rhinolophus lead us to reidentify the single specimen from Site 13, reported as R. subrufus by Heaney (1984), as R. inops. We netted two additional males in disturbed lowland forest at 1000 m (Site 4). Cranial and external measurements (Table 5) are slightly larger in most dimensions than those of the series from Biliran, Leyte (Rickart et al, 1993), and Mindanao (Mt. Kitanglad; Heaney et al, unpubl. data); we suspect that regional morphs corresponding to the Pleistocene islands (Heaney, 1986) are present. HEANEY ET AL: THE MAMMALS OF CAMIGUIN ISLAND 37


Specimens Examined — Total 3. Site 7 (2 fmnh); Site 13 (1 dmnh). Order Primates Family Cercopithecidae — Monkeys Macaca fascicularis (Raffles, 1821) The long-tailed macaque occurs from Burma to Timor and is found throughout the Philippines (Fooden, 1995; Heaney et al, 1998). It is found in agricultural areas near forest, in second-growth and secondary forest, as well as primary forest from sea level to at least 1800 m in lowland and montane forest (Goodman & Ingle, 1993; Rickart et al, 1993; Heaney et al, 1999). We did not collect any specimens, but in March 1995 we obtained the skull of an adult individual from a hunter who had killed the macaque in a forested area above Mahinog a few years earlier. Also in March 1995, a local hunter who was a member of our team saw two macaques a short distance from our camp at 1100m elevation (Site 7) in primary montane forest. He reported that the macaques were fairly common in what remained of lowland primary forest and good secondary forest at that time and were actively hunted. Specimens Examined — Total 1. Above Mahinog (1 fmnh). Order Rodentia Family Muridae — Rats and Mice Apomys camiguinensis Heaney and Tabaranza, 2006 A previously unknown species of Apomys (Musser, 1982) was trapped on Camiguin in May 1994 and March 1995 (Heaney & Tabaranza, 1997) and described in this volume (Heaney & Tabaranza, 2006b). The Camiguin forest mouse is moderately large for the genus, averaging about 40 g. It is marked by a number of subtle morphological characters, and molecular data show it to be most closely related to Apomys hylocoetes and A. insignis from Mindanao and to an unnamed species from Leyte and Biliran (Steppan et al, 2003; Heaney & Tabaranza, 2005b). This species was common (Table 1, Fig. 1) in disturbed lowland forest at 1000 m (Site 4) and in primary montane forest at 1275 m (Site 7) and relatively uncommon in primary mossy forest at 1200-1400 m (Site 6). It was not found in heavily disturbed lowland agricultural land (Site 3, elev. 150 m), lower montane forest (Site 4, elev. 1000-1300 m), or lower mossy forest (Site 8, elev. 1475 m), but there was limited sampling at all of these (Table 1). Most specimens were taken from traps placed in primary montane forest underneath root tangles and beneath fallen and rotten logs on a moderate slope. All individuals were captured at night. A pregnant female taken on 29 May 1994 weighed 405 g and had a single embryo (CRL 30 mm). Adult females with large mammae but not pregnant weighed 39 21 g (range 37542 g, N 5); two of these, taken 23 May 1994 and 17 March 1995, each had two placental scars. Nulliparous females (with small mammae) were lighter and had an average weight of 35 g (range 34-36). Females have two pair of inguinal mammae. These data indicate a litter size between one and two. Males with scrotal testes had a mean weight of 413 47 g (range 36548 g, N 11); testis size ranged from 15 X 16 mm to 8 X 12 mm. Males with abdominal testes weighed 330 50 g (range 28-34 g, N 5) and had testis size of 5 X 8 mm. Specimens Examined — Total 25. Site 4 (14 ms-iit), Site 6 (2 su-iit); Site 7 (9 fmnh). Bullimus gamay Rickart, Heaney and Tabaranza, 2002 A previously unknown species of Bullimus, recently described as B. gamay (Rickart et al, 2002), was captured on Camiguin in May 1994 and March 1995. The Camiguin forest rat was incorrectly assigned to Tarsomys by Heaney and Tabaranza (1997) and was designated "Bullimus sp. A" by Heaney et al. (1998). It was trapped from 900 to 1475 m (Fig. 1, Table 1) but not in heavily disturbed lowland agricultural land from sea level to 300 m (Site 3). Most specimens were taken from traps placed in runways beneath root tangles and rotten logs or near large rocks. This species was most common in primary montane forest at 1275 m (Site 7). It was less common in disturbed lower montane forest at 1000-1300 m (Site 5) and in primary mossy forest at 12001400 m (Site 6). It was rare in secondary lowland forest at 1000 m (Site 4) and in lower mossy forest at 1475 m (Site 8). Bullimus bagobus occurs in similar habitat on Mindanao but also occurs at lower elevation where good forest persists (Heaney, 2001; Heaney et al, unpubl. data); this suggest that B. gamay should also be sought at lower elevations on Camiguin. Parous females with large mammae weighed an average of 370 g (range 360-390 g, N 3), while nulliparous young adult females with small HEANEY ET AL: THE MAMMALS OF CAMIGUIN ISLAND 39


mammae were lighter, having a mean weight of 276 537 g (range 210-345 g, N 5). Adult males with scrotal testes had an average weight of 402 g (range 240-500 g; N 3) and had testis size ranging from 21 x 30 mm to 27 X 35 mm). Among the nine individuals taken in May 1994, the smallest was a nulliparous young female weighing 210 g, while among the 11 individuals captured in March 1995, two were juveniles, a male weighing 120 g and a female weighing 125 g. No pregnant females were taken in May 1994 and March 1995, but a lactating female (360 g) was taken on 18 March 1995, and a female taken on 29 May 1994 had three placental scars. This suggests that females give birth in February and March. Three females had three pairs of mammae and one had four pairs, indicating that litter size most likely does not exceed three. Specimens Examined — Total 20. Site 4 (4 msu-iit); Site 5 (2 ms-iit); Site 6 (3 su-iit); Site 7 (10 fmnh); Site 8 (1 fmnh). Crunomys melanius Thomas, 1907 The Mindanao shrew-mouse was previously known only from Leyte and Mindanao from near sea level to 1550 m, apparently in primary lowland rain forest; it is rare in collections (Musser & Heaney, 1992; Heaney et al, 1998; unpubl. data; Rickart et al, 1998). On Camiguin, we found it to be uncommon in heavily disturbed lowland agricultural land at 1000 m (Site 4), in disturbed lower montane forest at 1200 m (Site 5), and in primary montane forest at 1275 m (Site 7; Table 1, Fig. 1). It was trapped beneath rotten logs and wood tangles in an area with few dead leaves and little moss. It occurs down to sea level in good forest on Mindanao (Heaney et al, 1998; Heaney, 2001) and should also be sought at lower elevations on Camiguin. Two of the five specimens we captured were females. An adult female taken on 17 March 1995, with enlarged mammae and weighing 71 g, had recently given birth; it had three placental scars (one left, two right), and the uterus was swollen. The other female (625 g), probably nulliparous, had small mammae. Two adult males (60 g and 72 g) had scrotal testes (15 X 8 mm); the third male, taken on 24 May 1994, was a small juvenile (28 g) with abdominal testes. These scanty data imply that March, April, and May are months of reproductive activity. Females have two pairs of mammae, indicating that litter size is small. Cranial and external measurements (Table 6) are similar to but slightly larger than those from Leyte and Mindanao (Kitanglad Range and Mt. Apo; Rickart et al, 1993, 1998; Heaney et al, unpubl. data). Specimens Examined — Total 5. Site 4 (2 msu-iit); Site 5 (1 ms-iit); Site 7 (2 fmnh). Rattus everetti (Gunther, 1879) The common Philippine forest rat is a widespread endemic species (excluding the Babuyan Batanes, Palawan, and Sulu groups) found in primary and disturbed lowland, montane, and mossy forest from sea level to 2400 m (Heaney et al, 1991, 1998, 1999; Musser & Heaney, 1992; Rickart et al, 1993). During 1994-1995, we found this 250-350 g rodent to be fairly common at all sampling sites from disturbed lowland forest at 1000 m, through montane forest to mossy forest at 1475 m; specimens from the 1960s were taken at 500 m (Table 1, Fig. 1); on Mindanao, it occurs down to sea level (Heaney, 2001; Heaney et al, unpubl. data). It was trapped under root tangles and beneath fallen logs, in hollow spaces beneath live trees, and near mounds of soil and rotting leaves. Five parous adult females from May 1994 and March 1995 had a mean weight of 271 288 (range 240-300 g, N 5). During the same months, five nulliparous females (with adult pelage) weighed 188 214 g (range 160215 g), and four females with juvenile pelage weighed 116 310 g (range 77-150 g). Females have four pairs of mammae, indicating moderate litter size. None were pregnant, and we were unable to count placental scars; previous studies also produced little data on litter size (Heaney et al, 1999, unpubl. data). Males with scrotal testes, recorded in May 1994 and March 1995, had a mean weight of 288 698 g (range 210 375 g, N 4), and the largest males had testes size of 33 X 52 mm and 30 X 50 mm. Three males with abdominal testes, during the same months, weighed an average of 207 g (range 160-295 g). Cranial and external measurements (Table 6) are comparable to those from Biliran, Leyte, and Maripipi but slightly smaller than series from Dinagat, Mindanao (Kitanglad Range), and Siargao (Heaney & Rabor, 1982; Heaney et al, 1991, 1999, unpubl; Rickart et al, 1993). Specimens Examined— Total 28. Site 4 (6 msu-iit); Site 5 (2 ms-iit); Site 6 (3 su-iit); Site 7 (8 40 FIELDIANA: ZOOLOGY


fmnh); Site 8 (2 fmnh); Site 1 1 (4 dmnh); Site 12 (1 dmnh); Site 13 (2 dmnii). Ruins exulans (Peale, 1848) Known as the Polynesian rat or the spiny rice-field rat, R. exulans is not native to the Philippines; through mostly accidental dispersal by humans, it occurs from Bangladesh to Easter Island and throughout the Philippines, especially as a pest in agricultural areas. On Camiguin, we found this small (60-75 g) rat to be moderately common in heavily disturbed lowland agricultural land at ca. 150 m (Site 3) and in primary montane forest at 1275 m (Site 7) but not at other sites. Records from the 1960s also show it to be present in mossy forest at ca. 1500 m. This distribution is similar to that found on Negros, where it occurs in high-density agricultural areas and in mossy forest at an elevation of 15001650 m (Heideman et al, 1987; Heaney et al, 1989). Both Camiguin and Negros have very few native rodents, and this may influence the ability of nonnative species to invade the forest since the non-native are absent or very rare in mature forest on species-rich islands such as Leyte, Luzon, and Mindanao (Heaney et al, 1989, 1999. unpubl. data; Rickart et al, 1993). In March 1995, one nulliparous female weighing 41 g was trapped. Two adult males (71-76 g) with scrotal testes and two young adults (4961 g) with scrotal testes were trapped at the same time. They typically become reproductive at a young age and have many large litters each year (Barbehenn et al, 1973). Cranial and external measurements (Table 6) are comparable to those of Dinagat, Leyte, and Mindanao (Mt. Kitanglad), but slight variations in most dimensions are discernible (Heaney & Rabor, 1982; Rickart et al, 1993; Heaney et al, unpubl. data). Specimens Examined — Total 15. Site 3 (2 msu-iit); Site 7 (5 fmnh); Site 12 (1 dmnh); Site 13 (7 DMNH). Rattus tanezumi Temminck, 1844 Previously known in the Philippines as Rattus rattus and Rattus mindanensis, the Oriental house rat is a widespread nonnative rodent in the Philippines; it occurs from Afghanistan to Indomalaya. New Guinea, and Micronesia (Musser & Carleton, 1993). This 140 200-g rat is most often found as a pest in urban and agricultural areas but can be common in disturbed forest up to 1800 m (Heideman et al, 1987; Heaney et al, 1989, 1991, 1998, unpubl. data; Rickart et al, 1993). On Camiguin, we found R. tanezwni to be very abundant in heavily disturbed agricultural land at 150 m (Site 3; Table 1). It was less common in secondary lowland forest at 1000 m and in disturbed lower montane forest at 1000 m (Site 4) and 1200 m (Site 5), and a single individual was trapped in primary mossy forest at 1400 m (Site 6). Four nestling juveniles were found in a new slash-and-burn clearing near Site 7, but no R. tanezwni were caught in adjacent mature forest. Taken together with records from the 1960s, these data indicate that R. tanezwni occurs from sea level to the highest peaks on Camiguin, although it tends to be most common at lower elevations and in disturbed habitats. Pregnant females had a mean weight of 126 576 g (range 63-200 g, N 4). Females with large mammae but not pregnant had a mean weight of 151 167 g (range 130-169 g, N 9); three of these had placental scars (three, four, and eight scars). Nulliparous females had a mean weight of 93 398 g (range 67-160 g). Males with scrotal testes had a mean weight of 1694 227 g (range 141-199 g, N 8); a subadult with scrotal testes weighed 105 g. These data indicate large litter size (up to eight) and very early reproduction, as is typical for the species (Barbehenn et al, 1973). External and cranial measurements (Table 6) are comparable to those of Biliran, Leyte, Maripipi, and Mindanao, with only slight variation. Specimens Examined — Total 69. Site 3 (21 ms-iit); Site 5 (4 su-iit); Site 6 (1 su-iit); Site 7 (4 fmnh); Site 9 (2 dmnh); Site 10 (6 dmnh); Site 11 (3 dmnh); Site 12 (6 dmnh); Site 13 (22 dmnh). Order Carnivora Family Viverridae — Civets Paradoxurus hermaphroditus Jourdan, 1837 The common palm civet occurs from Sri Lanka to Hainan and the Lesser Sunda Islands and is widespread in the Philippines (Heaney et al, 1998). It has been recorded in agricultural and forested areas from sea level up to at least 2400 m (Heaney et al, 1991, 1999, unpubl. data). We did not capture any palm civets, but in March 1995 we were given a mandible from a specimen taken by local hunters in the mountains above Mahinog, thus confirming the earlier report by Gray (1843). A local hunter identified P. hermaphroditus from pictures as being common on Camiguin. 42 FIELDIANA: ZOOLOGY


Specimens Examined — Total 11 fmnh). Viverra tangalunga Gray, 1832 The Malay civet (or tangalung) is found from peninsular Malaysia to Sulawesi and is widespread in the Philippines (Heaney et al, 1998). It has been found in primary and secondary lowland, montane, and mossy forest from sea level to at least 1700 m (Rickart et al, 1993; Heaney et al, 1999). We did not obtain any on Camiguin, but a local hunter identified Viverra tangalunga from photographs as being present on Camiguin; we tentatively accept this as a valid record. Order Artiodactyla Family Suidae — Pigs Sus philippensis Nehring, 1 886 The Philippine warty pig is a Philippine endemic that occurs in the Greater Luzon, Greater Mindanao, and Mindoro faunal regions; its numbers are declining (Oliver, 1992, 1999). It formerly was abundant from sea level to at least 2800 m in all habitats; now it is common only in remote forests (Heaney et al, 1991, 1999, unpubl. data; Oliver, 1992, 1999; Garcia & Deocampo, 1997). On Camiguin in March 1995, we observed hoof marks of wild pigs from disturbed lowland forest at 1 000 m up to primary montane forest at 1275 m elevation (Site 7). Near the sampling site at 1475 m (Site 8), we saw an active pig nest, and scattered pig trails were in clear evidence. A local hunter said that they were commonly hunted and often sold in a small local market at Owakan, Mahinog Municipality, but not in coastal cities. We were given two adult mandibles by hunters from pigs captured in forest in the early 1 990s in the mountains above Mahinog. Specimens Examined — Total 2. Above Mahinog (2 fmnh). Analysis and Discussion Adequacy of Sampling: What Is Present and What Is Absent? Before interpreting field data, it is necessary to evaluate the extent to which they are complete and reliable. In doing so here, we follow the procedures used by Heaney et al. (1989, 1991, 1999) and Rickart et al. (1993). Fruit Bats — Because pteropodid bats lack sonar systems, they are easily captured in mist nets. The 1960s surveys, which focused on birds, yielded many fruit bat specimens (Heaney, 1984), most of them almost certainly from mist nets. Those efforts obtained six species of fruit bats (Fig. 2). Our netting in 1992-1994 yielded no additional species but was not extensive. We consider it quite possible that some additional species may be present, especially Eonycteris spelaea and Rousettus amplexicaudatus, since both occur very widely in the Philippines (Heaney et al, 1998), and possibly the large Acerodon jubatus and Pteropus vampyrus, which fly high above the canopy and so are difficult to capture. However, our sampling was sufficient that we consider it quite unlikely that the species that are abundant and easily captured on Mindanao and associated islands (Heaney et al, 1989, unpubl. data; Rickart et al, 1993) are present on Camiguin, especially Haplonycteris fischeri and Ptenochirus minor and probably Megaerops ecaudatus. Suitable habitat for the high-elevation specialist Alionycteris paucidentata is very limited in area on Camiguin, if present at all; although we did limited netting in that habitat, we think it very unlikely that this species is present. Sampling along the elevation gradient probably has produced a partial picture of variation in species richness (Fig. 2) but as noted may be incomplete because our netting in 1992-1995 was not extensive (Table 3). Insectivorous Bats — Insectivorous bats use sophisticated sonar systems to navigate. They are thus difficult to capture in mist nets, which were our only means of capturing them; further, our mist-netting efforts in 1992-1995 were limited in extent (see Heaney & Tabaranza, 2006a; Table 3). For this reason, we believe that our sample of six species is likely to be quite incomplete and is not usable for estimates of species richness, as is often the case (Heaney et al, 2002). Our data contribute, however, to general knowledge of the natural history of these poorly known animals. Nonvolant Small Mammals — Shrews, rodents, and other small mammals that can be captured in traps were poorly sampled by the 1 960s field teams, which focused their efforts on birds (and secondarily on fruit bats). During 1994 and 1995, we accumulated 2783 trap-nights at six sites that included all major habitat types on the island, yielding 188 captures (Table 1). A species-accumulation curve for these data (Fig. 3) shows that the number of species recorded for HEANEY ET AL: THE MAMMALS OF CAMIGUIN ISLAND


No. Species 200 400 600 800 1000 1200 1400 1600 1800 2000 2200 2400 2600 2800 No. Trap-nights Fig. 3. Species accumulation curve for nonvolant small mammals on Camiguin Island. The points shown are for Sites 1 and 3 to 6. as described in the text. the island reached a peak of eight species (including five nonnative species) by the end of sampling at the second site (1055 trap-nights) and showed no further gain as we approached the total of nearly 2800 trap-nights. Local hunters either were unfamiliar with squirrels or knew them from Mindanao; given their conspicuousness (Heaney et al, unpubl. data), they are very unlikely to be present. We conclude that the island is unlikely to support other species of small mammals that are present on Mindanao, including gymnures (Podogymnura), tree shrews (Urogale), squirrels, or murid rodents (eg, Batomys, Limnomys, and Tarsomys). Further, the sampling along the elevational gradients was sufficiently intense that we are likely to have an accurate assessment of the pattern of species richness above about 800 m, although it must be noted that extensive habitat destruction at lower elevations may have had an extensive impact on the current patterns (discussed below). Medium and Large Mammals — We collected information only on the presence and absence of large mammals and a mixed set of medium-sized species (none of which are readily captured in traps such as the ones we used), not on detailed habitat use. We observed carefully during all our time in the field for any evidence of additional species, and we interviewed many residents of the islands and showed them photos of potential species; some of these residents are active hunters. Neither we nor our informants had any evidence for tarsiers, flying lemurs, or deer on the island, though many people knew of deer and tarsiers on Mindanao, and a few knew of flying squirrels and flying lemurs. We consider it unlikely that species other than those listed here are present given the voucher specimens we obtained and the concordance between our specimens and the information received from local hunters. Thus, we conclude that deer are absent, along with the smaller but conspicuous species such as tarsiers, squirrels, and flying lemurs. Elevational Patterns Though our limited netting prevents us from commenting on the patterns of diversity and abundance of bats along the elevational gradient on Camiguin, our trapping data (Table 1) provide a reasonably complete picture of the general patterns among nonvolant species. The highest density we encountered was at Site 3, in agricultural land at ca. 150 m. However, these were all nonnative rats and a shrew, representing only three species, a pattern that we have often seen (eg, Heaney et al, 1989; Rickart et al, 1991). The five native small mammal species were all present (or suspect to be present) from 1000 to 1300 m and declined in species richness to two species near the peak of Mt. Timpoong at Site 8 (Table 1). Density (measured as the number captured per 100 trap-nights) more than doubled from 1000 to 1275 m, then fell to a much lower level near the peak at 1475 m. However, much of the change in density was driven by the remarkably large numbers of Suncus murium at the upper elevations; they were not taken at 1000 or 1200 m (though they were taken at 150 m), were fairly common at 1300 and 1475 m, and were exceptionally abundant at 1275 m, where we often saw and heard them during the day. These patterns for species richness, though limited in extent, are similar to those from elsewhere in the Philippines, where species 44 FIELDIANA: ZOOLOGY


Incisors Condylobasal length. Condylocanine length. Fig. 5. Bat skull with teeth and cranial measurements indicated. age and sex. It should then be identified to family with the Key to Families, and then to species with the key to that family. Next, all external measurements should be taken and matched against those in the tables of measurements; if the measurements do not match, the identification is questionable and the process should be begun again from the first step. Finally, if a skull is available, it should be visually compared with Figures I960 to see if it matches the features of that species or genus. Determination of Age and Sex The age of bats can be estimated by the degree of ossification of the joints in the digits of the wing (Anthony, 1988). In juvenile bats, the joints have cartilaginous discs where growth takes place. When a light is shown through the wing of a live bat, the bands of cartilage at the joints appear partly translucent (fig. 6B). In adult bats, the bones are fully ossified and the joints appear opaque (fig. 6 A). The shape of the joints of the digits also differs between juvenile and adult bats; this is especially useful in determining the age of animals prepared as stuffed skins or fluid-preserved specimens. In juvenile bats the joints are swollen and tapered (fig. 6B), whereas in adult bats the joints are knobby and more distinct from the bone shaft (fig. 6A). Bats can be sexed by examination of the external genitalia. Males have a conspicuous penis (except B Fig. 6. Joints in digits of wing. Stippled areas represent bone and open areas represent cartilage. A, adult; B, juvenile. INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS


mm Fig. 7. Tails of representatives of Philippine bat families. A, Pteropodidae (Rousettus); B, Megadermatidae (Megaderma); C, Vespertilionidae (Miniopterus); D, Rhinolophidae (Rhinolophus); E, Molossidae (Chaerephon); F, Emballonuridae (Taphozous). Family Pteropodidae: Fruit Bats All fruit bats are characterized by dog-like heads (fig. 8A), with eyes that are proportionately larger than those of most other bats; the large eyes are associated with their dependence on vision for orientation and their lack of echolocation (sonar) systems (except for the very simple and limited one used by Rousettus). They do not have the elaborate Fig. 8. Heads of representatives of Philippine bat families (not to same scale). A. Pteropodidae (Rousettus). Note that the ear margin is continuous, neither tragus nor noseleaf is present, the eyes are large, and the face is dog-like. B. Megadermatidae (Megaderma). Note that a noseleaf is present, the tragus is forked, and the large ears are connected across the top of the forehead. C. Rhinolophidae (Rhinolophus). Note the elaborate noseleaf, the well-developed FIELDIANA: ZOOLOGY


p y YJWJ» X antitragus, and the absence of a tragus. D. Emballonuridae (Taphozous). Note that there is no noseleaf and that a tragus is present. E. Molossidae (Chaerephon). Note that there is no noseleaf and that the ears are thick and fleshy. F. Vespertilionidae (Miniopterus). Note that there is no noseleaf and that a tragus is present. INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS


Fig. 10. Dorsal views of fruit bat muzzles (not to same scale). A, Macroglossus; B, Haplonycteris; C, Nyctimene; D, Rousettus; E, Cynopterus. 1 1A); teeth, except canines, greatly reduced; forearm 41-45 mm; skull as in Figure 26 Macroglossus minimus 16. Muzzle short and broad; nostrils tubular (as in fig. 10B); wing membrane attaches on side of foot (on or above outermost toe; fig. 1 1C); teeth robust; forearm 43-53 mm 17 17. Four upper and two lower incisors (fig. 9D); forearm 45-52 mm Megaerops wetmorei 17. Two upper and two lower incisors (as in fig. 9 A); forearm 43-53 mm 18 18. Band of pale fur along dorsal surface of forearm; interfemoral membrane present; forearm 46-53 mm; skull as in Figure 24 Haplonycteris fischeri 18. No band of pale fur along dorsal surface of forearm; no interfemoral membrane; forearm 45-50 mm Alionycteris paucidentata 19. Ears and skin on dorsal surface of bones of wings with prominent pale yellow spots; dark stripe along most of dorsal midline; two upper and no lower incisors (fig. 9Q; nostrils elongated into tubes about 2-3 mm long (fig. IOC); forearm 71-79 mm; skull as in Figure 27 Nyctimene rabori 19. Ears and wings without yellow spots; no stripe along dorsal midline; four upper incisors and at least two lower incisors; nostrils not elongated into tubes (although when viewed from above they may appear slightly tubular); forearm 45-92 mm 20 20. Muzzle moderately long and tapered (fig. 10D); anterior surface of upper canines with vertical groove (not always prominent); wing membrane terminates above gap between outermost toe and second toe from outside (fig. 1 1 B); forearm 80-92 mm; skull as in Figure 32 Rousettus amplexicaudatus 20. Muzzle short and broad (fig. 10E); anterior surface of upper canines smooth, not grooved; wing membrane terminates either on outermost toe (fig. 1 1C) or above gap between outermost toe and second toe from outside (fig. 1 1 B); forearm 45-92 mm 21 21. Four upper and two lower incisors (outer pair of upper incisors much smaller than inner pair; fig. 9D) 22 21. Four upper and four lower incisors (fig. 9B) 23 22. Forearm 76-90 mm; skull as in Figure 28 Ptenochirus jagori INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 13


The Birds of Sibuyan Island, Romblon Province, Philippines, with Particular Reference to Elevational Distribution and Biogeographic Affinities Steven M. Goodman David E. Willard Pedro C. Gonzales Abstract Sibuyan Island is a relatively small, isolated island measuring at its widest points 24 km north to south and 28 km east to west, rising to 2050 m above sea level, with a total land area of 233 km. The nearest major island masses are Luzon (to the northeast), Mindoro (to the northwest), and Panay (to the south). A considerable portion of the natural habitat, including some lowland forest, remains intact. Sibuyan is oceanic in origin and during the middle to late Pleistocene was not attached to any other island mass. Most of the small islands in the Philippines for which we have ornithological information were connected to larger island groups during the Pleistocene, making an analysis of the avifauna of Sibuyan, and its faunistic relationships to other surrounding larger islands, important in understanding patterns of dispersal across water barriers, and providing an interesting contrast to the avifaunas on Pleistocene land-bridge islands. A total of 1 30 bird species have been recorded on Sibuyan: 102 known or presumed residents and 29 migrants or boreal winter visitors. Historical records combined with the results of our 1989 and 1992 field trips show 93 resident species occurring between and 100 m, 24 at 325 50 m, 19 at 725 50 m, and 10 at 1325 50 m. Although little has been published previously on the birds of Sibuyan, the number of collections from the island is remarkable, spanning almost 100 years of ornithological exploration. Several collectors described the habitats they found on the island during their visits. This information provides a window into the extinction and colonization rates of birds as a function of time and habitat degradation. The cutting of the lowland forests for lumber or conversion into agricultural areas has had a major impact on the avifauna. Three or four species of large pigeons have likely been extirpated on Sibuyan. Five species of open country birds have colonized the island in the past few decades. The resident bird species found on Sibuyan are closest to those on the islands of Mindanao, Luzon, Mindoro, and Panay. This group of islands has an avifauna that is very distinct from Palawan Island's. Sibuyan is one of the last small islets in the Philippines with significant large tracts of forest. Its relative remoteness and lack of extreme human pressure on the remaining natural resources, combined with a biologically diverse environment, make it an ideal area for a reserve. The Department of Environment and Natural Resources, Manila, is currently considering a plan to place a portion of the island in the national protected area system. Introduction tially or totally cleared of their lowland forests. Fewer than 10% of these islands have had avifauthe Philippines comprise over 7,000 islands and nal surveys, and the percentage of well-surveyed islets, the vast majority of which have been par-islands is even smaller. To date, virtually all of the FIELDIANA: ZOOLOGY, N.S, NO. 82, NOVEMBER 30, 1995, PP. 1-57 1


Fig. 1 Position of Sibuyan Island relative to other islands in south-central Philippines. Sibuyan is located just below center on the map. The islets between Mindoro and Panay are numbered as 1— Semirara, 2— Sibay, 3— Caluya, and between Panay and Tablas as 4— Carabao, 5— Boracay. studies of birds occurring on smaller islands have been conducted on those connected by land bridges to much larger islands during the late Pleistocene (eg, Catanduanes 1,823 km, connected to greater Luzon] — Gonzales, 1983; Marinduque 1 727 km, connected to greater Luzon] dupont, 1972b; Ticao 362 km, connected to greater Panay-negros— dupont, 1972a; Dinagat 668 km, connected to greater Mindanao] dupont and Rabor, 1973; and Siargao 455 km, connected to greater Mindanao— dupont and Rabor, 1973). There is no published study in the Philippines of the birds found on a small island of oceanic origin and surrounded by deep water. Information on the vertebrates of isolated oceanic islands, with relatively intact forest, helps address biogeographic questions of dispersal patterns and origins of taxa colonizing across water rather than across land (eg, Heaney et al, 1989, 1991). In the late 1980s, Sibuyan was identified as an important site because it is oceanic in origin, is surrounded by deep water, is relatively isolated, and has considerable elevational relief. Sibuyan is separated from Romblon by a 105-km-wide channel that is at least 180 m deep (Fig. 1). Romblon lies 95 km east of Tablas, and these two islands are separated by an ocean depth of at least 55 m. The southern edge of Tablas is about 43 km north of Panay Island, the nearest large island to the south, and the small islets of Carabao and Boracay lie in between. The ocean floor depth between these islands varies: Boracay and Panay— 20 m; Carabao and Panay— 90 m; and Carabao and Tablas— at least 350 m (Philippine Coast Pilot, 1 968; Defense Mapping Agency Hydrographic Center, 1977, 1978). Carabao and Tablas are separated by a channel less than 5 km wide. Approximately 1 7,000 years ago, during the last FIELDIANA: ZOOLOGY


Table 2. Continued. Brackets indicate that the elevation is inferred. Habitat definitions based mostly on information from Sibuyan Island and supplemented by dupont (1971) and Dickinson et al. (1991): A aquatic, including rice fields; O open or heavily disturbed areas; and F forest or forest edge. (accrued net-days in parentheses) to 30 m, 028 bird per net-day (90); 325 m, 028 bird per net-day (151); 725 m, 032 bird per net-day (140); 1325 m, 018 bird per net-day (78); and 1525 m, 013 bird per net-day (36). Thus, general capture rates were more or less constant through 725 m; above this elevation the numbers declined substantially. Migrants on Sibuyan Island— Of the 1 32 species recorded on Sibuyan, 29 (22) are not known to breed on the island. This includes migrants that spend the boreal winter on the island and return in the spring to mainland Asian or Japanese breeding grounds, and those that pass through the Philippines in the autumn and spring while on migration. Of the 29 migrants recorded on the island, GOODMAN ET AL: BIRDS OF SIBUYAN ISLAND, PHILIPPINES 41


Mcgregor's travel itineraries, we know that they did not visit the higher slopes of Mt. Guitinguitin. They presumably hired local hunters to gather birds from a variety of localities and habitats. Given the broad elevational range of several of these pigeons elsewhere in the Philippines and the fact that intact forest remains on Sibuyan within this range, the disappearance of these birds cannot be explained simply by the destruction of the lowland forest. Further, there is no evidence that pigeon hunters have ever visited remote upland areas of the island. Large frugivorous pigeons in the Philippines are known to have elevational movements associated with the availability of fruits, often in large flocks and over considerable elevational ranges (Rabor, 1977; Miranda, 1987). It is conceivable that the pigeons that formerly occurred on Sibuyan depended seasonally on lowland fruit trees. The destruction of most of the lowland habitat, combined with locally intense human hunting pressure in the areas that pigeon flocks would have visited, may have led to their disappearance from he island. Cumulative records of the occurrence of Ducula poliocephala on Sibuyan may support the notion of interisland migration for this species. Mcgregor ( 1 905a) reported collecting this species on Sibuyan both in interior forest and near the sea. The only subsequent records of this vocal and sometimes conspicuous species on the island are of nine individuals taken at four different localities in late August and October 1971. The sporadic nature of these records points to erratic seasonal movements between islands. However, similar movements do not seem sufficient to explain the disappearance of the other two Ducula and the Columba pigeons from Sibuyan. The fact that early explorers found all of these species during short visits to the island and none was found after 1928, despite use of local hunters and more thorough exploration of all island habitats, indicates that these changes are real and not just a question of sampling or chance. Other species of forest-dwelling birds such as Megapodius cumingii and Tanygnathus lucionensis have serious declined on Sibuyan. Megapodius was found on Sibuyan by Bourns and Worcester (1894) and Mcgregor (1905a) but was not positively documented thereafter by other scientists visiting the island. In other areas of its range, this species often occurs in lowland and coastal forests (Delacour & Mayr, 1946; Dickinson et al, 1991), and it is often hunted or the nests robbed of eggs (Dickinson et al, 1991; SMG, pers. obs). In the 1 950s eggs of this species were seasonally available in the Magdiwang market (J. L. Tansiongco, pers. comm). In October 1991a bird hunter from Tampayan shot at least two Megapodius in the mangroves at Agnonok. Presumably, the combination of habitat change and human persecution has led to this species' rarity on the island. Tanygnathus was apparently more common at the turn of the 19th century than in recent years. An informant, about 50 years old and a resident of Jao-asan, mentioned that he had seen this species go from common to scarce within his lifetime. There are some anomalous cases of species known to occur in a variety of habitats, including those still present on Sibuyan, that were recorded by early ornithological expeditions to the island but not seen subsequently. These include Ardea purpurea, Egretta sacra, Dendrocygna arcuata, Falco peregrinus. Halcyon capensis, and Halcyon coromanda. In most cases these species were documented only once on the island, and the subsequent lack of records may be best explained by migration, rarity, or simply chance. With the conversion of much of the lowland forest through logging and agriculture, creating new habitats, a number of species that are tolerant of disturbance have colonized the island. One is Pycnonotus goiavier, which throughout its broad Southeast Asian range is regarded as "common in scrub, second growth, residential, and cultivated areas" (Dickinson et al, 1 99 1 p. 284). Until 1 954 it had not been recorded on Sibuyan. Since that period it has proliferated and is now one of the most common species in heavily disturbed areas below 75 m. Other examples include two species of open-country doves, possibly introduced to other Philippine islands by humans. Geopelia striata was first recorded on Sibuyan in 1972 and Streptopelia chinensis in 1989. Passer montanus, another species introduced into the Philippines, was first recorded on Sibuyan in 1989 at Magdiwang. In 1992 we found it in all of the large coastal villages around the island. Bubulcus ibis is first known from the island in 1972. Other recent additions to the Sibuyan list are simply the result of more intensive exploration of higher elevations (Turdus poliocephalus and Zosterops montana) and the discovery of species rare enough to have been missed or with secretive behavior (Ixobrychus sinensis, Porzana fusca, Amaurornis olivaceus, A. phoenicurus, and Cuculus fugax). In summary, the avifauna of Sibuyan has undergone some clear changes in the past 1 00 years. In several cases the recent addition of species tol-GOODMAN ET AL: BIRDS OF SIBUYAN ISLAND, PHILIPPINES 43


Table 4. Continued. Species Tablas Romblon Sibuyan Sources' Collocalia troglodytes Alcedo c. cyanopectus Ceyx lepidus margarethae Halcyon capensis gigantea Halcyon coromanda major Halcyon smyrnensis gularis Halcyon winchelli nesydrionetes Halcyon chloris collaris Merops viridis americanus Merops p. philippinus Eurystomus orientalis cyanocollis Megalaima haemacephala homochroa Dendrocopos maculatus menagei Pitta e. erythrogaster Pitta s. sordida Alauda gulgula wolfei Hirundo tahitica javanica Coracina striata mindorensis Lalage nigra chilensis Pycnonotus g. goiavier Hypsipetes siquijorensis cinereiceps Dicrurus hottentottus menagei Oriolus chinensis yamamurae Corvus macrorhynchos philippinus Copsychus saularis mindanensis Turdus poliocephalus subsp. Megalurus timoriensis tweeddalei Megalurus palustris forbesi Cisticola exilis semirufa Cyornis rufigastra philippinensis Rhipidura a javanica nigritorquis Rhipidura cyaniceps sauli Hypothymis coelestis rabori] Hypothymis a. azurea Terpsiphone cinnamomea unirufa Pachycephala homeyeri winchelli Anthus novaeseelandiae lugubris Artamus I. leucorhynchus Aplonis p. panayensis Sarcops calvus melanonotus Anthreptes malacensis chlorigaster Nectarinia sperata trochilus Nectarinia. jugularis Aethopyga siparaja magnifica Dicaeum aeruginosum striatissimum Dicaeum trigonostigma intermedium Dicaeum trigonostigma sibuyanicum Dicaeum trigonostigma cnecolaemum Dicaeum p. pygmaeum Zosterops montanus subsp. Passer montanus saturatus Lonchura leucogastra manueli Lonchura malacca jagori Total number of species 74 57 102 Sources for Tablas and Romblon records: 1— Bourns and Worcester (1894), 2— Mcgregor (1905a), 3— Baud (1976), 4— specimens) in amnh, 5 — specimens) in bmnh, 6— specimens) in fmnh, 7— specimens) in pnm, 8— specimens) in such, 9— specimens) in usnm, 10— Mcgregor (1906b), 1 1 Dickinson et al. (1991), 12— Mcgregor (1909), 13— Mcgregor (1910), 14— specimens) in cm, 15— specimens) in dmnh. See text for documentation of Sibuyan records. Names in bold are endemic forms. Records in brackets denote that the occurrence of a species on an island has not been verified by a specimen. GOODMAN ET AL: BIRDS OF SIBUYAN ISLAND, PHILIPPINES 47


ft 59 f? FIELDIANA ZOOLOGY op. 9 Published by CHICAGO NATURAL HISTORY MUSEUM Volume 39 September 18, 1961 No. 52 A New Race of Crow, Corvus enca, from the Philippines Austin L. Rand Chief Curator, Department of Zoology AND DIOSCORO S. Rabor Associate, Department of Zoology In the course of Dr. Rabor's field work in northeastern Luzon, in 1960, he collected two specimens of the crow, Corvus enca, which provide a substantial range extension of the species, and, as he recognized, represent an undescribed subspecies. For this we propose the name: Corvus enca sierramadrensis, new subspecies Type. — Chicago Natural History Museum No. 260,006, from San Pascua, Gonzaga, Cagayan Province, in Sierra Madre, Luzon. Adult female, collected April 20, 1960, by D. S. Rabor. Diagnosis. — Most like C. e. samarensis, from Samar, but bill shorter and more slender, culmen less arched; concealed bases of feathers of neck and forepart of body grayer; plumage generally more intensely black. Range. — Known only from Cagayan Province of extreme northeastern Luzon. Remarks. — The discovery of this species on Luzon has some zoo-geographical implications. Hitherto, its supposed absence there has suggested that some historical factor might have been responsible for its not reaching Luzon. Now that we know that it occurs in the three largest main islands of the Eastern Philippine Province (Luzon-Samar-mindanao), as well as the Mindoro District and the Palawan group (small and large islands), it would seem that the distribution pattern and the great variations in abundance in the Philippines Library of Congress Catalog Card Number: 61-17859 -. No927 577 IIE LIBRARY OFT NOV 27196 mez 1 TY OF IU


272 SPECIES BLANCOANAE 18 cm) long, while the pericarp of the fruit is described as fragile. The description conforms to Parashorea plicata, and I have not the slightest hesitation in adjusting the synonymy. The native name cited by Blanco, malaanonan, is valueless in interpreting the species, as it is a made up one, literally "false anonang," anonang = Cordia myxa Linn., and, as used to-day, is very loosely applied, although I have specimens of Parashorea plicata Brandis from Laguna Province, Luzon, bearing this name. Illustrative specimen from Mount Maquiling, Laguna Province, Luzon, comm. N, Catalan, June 20, 1914, with the Tagalog name malaanonang (Merrill: Species Blancoanae No. 1053). VATICA Linnaeus VATICA MANGACHAPOI Blanco Fl. Filip. (1837) 401 (sp. nov.) =V atica apteranthera Blanco op. cit. ed. 2 (1845) 281 (nom. nov.) ; ed. 3, 2 (1878) 156. Vatica sinensis Blanco op. cit. 401; 280; 156 (chinensis), non J. F. Gmel. ? VATICA MANGACHAPOI Blanco. The correctness of this interpretation of Vatica mangachapoi Blanco is very doubtful, but i have followed Vida, Brandis, and Foxworthy. It is to be noted that the name mangachapoi is apparently never applied to the species here as interpreted, but is used for Hopea acuminata Merr., H. pierrei Hance, and perhaps some other species. Vatica sinensis is said by Blanco to differ from V. mangachapoi in having anthers winged, so that probably the plant he described under this name was not the same as his V. aptheranthera (= V. mangachapoi). As to the anthesis, Vatica mangachapoi, as here interpreted, produces flowers from December to June in the provinces near Manila, while Blanco indicates May for both species discussed above. Foxworthy has placed here also Mocanera mangachapoi Blanco Fl. Filip. (1837) 450 (sp. nov.) = Dipterocarpus mangachapoi, and Blanco’s description of the leaves as “membranaceas” does not apply to S. balangeran. The identity of the species may later be determined from special collections in Bulacan and Rizal with reference to the native names cited by Blanco, and I suspect that the species is


No. 35) HANNA: WEST AMERICAN COS VS 11 43 Type locality: For fergusoni, Panama; for xanthicus, off Guaymas, Mexico. Range: Turtle Bay and Magdalena Bay (outer coast), Baja California and Gulf of California, Mexico, south to the Galapagos Islands and Mancora, Peru (D. L.frizzell). Collecting Stations: Mexico: Arena Bank, Baja California, (Sta. 136, D-11, 24, 27, 30 to 50 fms); Santa Inez Bay, Baja California, (Sta. 142, D-4, 40 to 50 fms; Sta. 147, D-2, 60 fms); Gorda Banks, Baja California, (Sta. 150, D-13,16, 25, 56 to 80 fms); off Pyramid Rock, Clarion Island, (Sta. 163, D-4, 50 fms); Chamela Bay, (Sta. 182, D-4, 16 fms)-Costa Rica: Port Parker, (1 to 90 fms); 14 miles SXE of Judas Pt, (Sta. 214, D-l, 2, 3, 4, 42 to 61 fms); Panama: Hannibal Bank, (Sta. 224, 35 to 40 fms). Above are New York Zoological Society Stations. Additional Calif. Acad. Sci. localities are Turtle Bay, Santa Maria Bay, Magdalena Bay and Cape San Lucas, Baja California; Clarion Island. It seems remarkable that this huge species, sometimes over five inches long, should have remained undiscovered until 1873, but a search of the literature has failed to disclose a name for either the white adult form or the colored juveniles. Dall suggested that C comes Hinds (1843, p. 256), might be the young of fergusoni although both Hinds (1844, p. 7), and Reeve (1843, pi. 13, fig. 64, Suppl. p. 4), came to believe their shell to be the young of C terebellum, while Tryon (1884, p. 80), after copying Reeve's figure, considered it to be C. terebra. De Barros e Cunha (1933, p. 183), followed Tryon and cited terebra from the Philippines and North Australia. Hinds stated explicitly that coelebs was found on a Fiji coral reef and his localities are usually trustworthy. Conus quercinus Hwass in Bruguiere (1792, p. 681), is very similar to the large fergusoni; specimens in the California Academy of Sciences labelled from Mauritius, can hardly be distinguished. Conus virgo Gmelin (1789, p. 3376); from the same region is also similar but is shaded with purple on the interior of the lower end of the aperture. An earlier name for the species seems to be Conus cingulum Martyn (1784, vol. 1, fig. 39). A magnificent series of growth stages has shown that C. xanthicus Dall is the young of fergusoni. Without such a series it would not be suspected that the bright yellow or orange small shells could possibly be the same as the huge white ones so familiar to beach collectors at Magdalena Bay and vicinity. For a long time we were puzzled that no small white ones ever turned up in any of the collections. It develops that when xanthicus reaches a length of 60 mm. the colored bands become very faint yet upon closer inspection it is found that they still persist in specimens which are


is explained by the high population of this fish off the coast of Norway and the use of purse spines, which accounted for 455,000 t. The Soviet Union has greatly intensified its fishing for mackerel in recent years, in the northwestern Pacific (to 80,000 t) and off West Africa (to 150,000 t) Doubtless, mackerel fishing is quite promising. The catch of Carangidae had increased by a factor of 16 by 1970 in comparison to the prewar level (1938), and there is every reason to assume the possibility of further increases. Thus, off the coast of Japan in some years the catch of this fish has even been artificially held down. Large accumulations of mackerel have recently been found off the Tasmanian coast. In the opinion of many scientists, it will be possible to catch as many as 1 million tons of pelagic fish per year near New Zealand, including many Carangidae. The greatest quantity of Japanese mackerel are caught in the East China Sea and off the coast of Japan (510-56010t), as well as in the Philippines (LOOLO t) Other species of Carangidae are caught off the west coast of Africa, where fishing is conducted primarily by ships of the USSR (ISOLO-t) Angola (170 10 t) and Spain (70 -lO-t). A lesser role in the sea fish catch (up to 1710" t) is played by the tuna, although due to the high value of the meat of this fish, tuna fishing is significantly more successful economically than fishing for most other types of fish. Therefore, the tuna catch develops from year to year, and at the present time is conducted in the open Pacific, Atlantic and Indian Oceans. Tuna inhabit the great water areas of the ocean, forming no dense accumulations, making it more difficult to catch them. However, information on the results of tuna fishing indicates that the trend is doubtless toward decreasing effectiveness (Table 12). For example, the catch of yellowfin and blue-finned tuna decreased from 8 fish per 100 hooks on a longline in 1950 to 2 in 1965. The reserves of large tuna are being caught with great intensity in almost all regions of the ocean, and the only hope for any significant development of tuna fishing lies in more intensive fishing of the smaller and stripe-bellied tuna. The overwhelming majority of tuna (50-60) is caught by Japan. Significantly smaller quantities are caught by the fisherman of the USA, France, Spain and Peru. The Soviet Union catches but a few thousands of tons of the valuable fish. When tuna is fished using longlines, swordfish and sharks are caught in large quantities. Sharks, rays and chimaera are also the objects of a specialized fishing industry. The annual catch of sharks-in recent years (1968-1975) has held steady at a level of about 400' 10 t, of rays at 90-140 -lO-t. This is much less than their actual reserves. The shark catch in the World Ocean could be at least doubled. In addition to the commercial fish mentioned above, the world catch includes many representatives of various families of demersal fish, caught during trawling and other types of bottom fishing: Pleuronectidae (see Table 8), sea perch, Ammodytes, etc. (Table 13). 374


SPEARFISHES OF THE CENTRAL PACIFIC 521 to a dark, slate gray on the back and to black on the first dorsal. We have seen no evidence of stripes and, according to Nakamura (1949), it never has them. Distribution in the Pacific According to Nakamura (1951), this pelagic species does not enter coastal or enclosed seas. Off Japan it occm-s south of 35° N. latitude and rather densely in the waters east of Formosa and the Philippines from November to January. Nowhere is it abundant enough to be of importance to the fishery. In our POFI fishing we have taken only the 8 specimens recorded in appendix table a; their distribution is indicated in figure 23. In the Hawaiian fishery it is one of the miscellaneous spearfishes that comprises only' a small fraction of the total spearfish catch. On the first six Japanese mothership expeditions to the vicinity of the Caroline Islands (Ego and Otsu, 1952) it was combined with the sailfish in the statistics, and on each of these trips the catch of the two species together averaged only from 02 to 07 per 100 hooks. Size This is the smallest of the spearfishes and, according to Nakamura (1949), attains a weight of only 44 pounds, but the POFI specimens which we have weighed from the central Pacific ranged from 18 to 51 pounds. Based on the data obtained from the Honolulu market by the Hawaiian Division of Fish and Game (table 3), the maximum weight found in 177 specimens was 114 pounds. However, the modal size was approximately 38 pounds. FIGURE 215. — Distribution uf POFI catches of shortiioso spearfish, Tetraplurus angustirostris. Fractions indicate stations at which catches were reported out of the total fished; decimals indicate average catch per 100 hooks per day.


FISH FROM THE HAWAIIAN, MARSHALL, CAROLINE AND PHILIPPINE ISLANDS, ALSO FROM FRENCH INDO CHINA AND MALACCA STRAITS No. 318 Philippine Wrasse, Labridae specie South China Seas Yacht ARA, January 23, 1929, Manilla, Philippine Islands 319 "Luau" Red Parrotfish, Callyodon miniatus Western Pacific Ocean Yacht ARA, December 14, 1928, Honolulu, Hawaiian Islands 320 Silver Bream, Seams sara Indo-pacific Waters Yacht ARA, February 11, 1929, Georgetown, Penang, Malay States 321 Square-tailed Sea Perch, Plectropomus truncatus South China Seas Yacht ARA, January 23, 1929, Manilla, Philippine Islands 322 Blue Lined Butterfly Fish, Chaetodon fremblii Pacific Ocean Yacht ARA, December 14, 1928, Honolulu, Hawaiian Islands 323 Godeffroy's Wrasse, Anampes godcffroyi Malacca Straits Yacht ARA, February 8, 1929, Singapore, Malay States 324 Unicorn Surgeon Fish, Acanthurus unicornis Hawaiian Islands, Tropical Pacific Ocean Yacht ARA, December 14, 1928, Honolulu, Hawaiian Islands 325 Short-Snout Unicorn Fish, Acanthurus brevirostris Western Pacific Ocean Yacht ARA, December 14, 1928, Honolulu, Hawaiian Islands 326 Indian Conger Eel, Muraenesox cinereus South China Seas Yacht ARA, January 23, 1929, Manilla Philippine Islands 32


No. 368 Great Dog Herring, Chirocentrus dorab Malacca Straits, Red Sea. Indian Ocean Yacht ARA, February 11, 1929, Georgetown, Penang 369 Red Rockfish, Heucohnus rufescens Western Pacific Ocean Yacht ARA, December 15, 1928, Kenohe Bay, Hawaiian Islands 370 Black Harvest Fish, Stromatnis niger East Indian Seas Yacht ARA, February 11, 1929, Georgetown, Penang 371 Spotted Butterfly Fish, Scatophagus argus Indian Ocean to Australia Yacht ARA, January 23, 1929, Manilla, Philippine Islands 372 Purple Flying Fish, Cypsuiinis shuns Tropical seas about Hawaii Yacht ARA, December 11, 1928, Hilo, Hawaii, Hawaiian Islands 373 Thread Herring, Pellona filigera East Indian Seas Yacht ARA, February 11, 1929, Georgetown, Penang, Malay States 374 Lined Bark-scaled Wrasse, Lepidaplois lunulatus Western Pacific Ocean Yacht ARA, December 14, 1929, Honolulu, Hawaiian Islands 375 Indian Carp, Cyprinidae specie Saigon River, Indo-china Yacht ARA, January 30, 1929, Saigon, Indo-china 376 Square-tailed Sea Perch, Plectropomus truncafits South China Seas Yacht ARA, January 23, 1929, Manilla, Philippine Islands 377 Archer Fish, Toxotes jaculator Found throughout East Indies Yacht ARA, January 30, 1929, Saigon, Indo-china 378 Poor Halfbeak, Hemiramphus depauperatus Tropical Pacific Seas Yacht ARA, December 11, 1928, Hilo, Hawaii, Hawaiian Islands 37


PONYFISH LIGHT ORGAN SYMBIOSIS 411 that cover the light organ, from the internal location of the light organ, and from several unique anatomical adaptations of the gas bladder, body musculature, and skin of the fish (Haneda, 1940; Mccall-ngai, 1983a, b; Mcfall-ngai and Dunlap, 1983, 1984; Dunlap and Mcfall-ngai, 1984). The internal, poorly vascularized ponyfish light organ overlies the esophagus just anterior to the stomach and interfaces with the oxygen-rich gas bladder across a thin, window-like membrane that is permeable to oxygen (Mcfall-ngai, 1983a, b). Within the light organ, the dense, pure culture of P. leiognathi (Boisvert et ai, 1967; Hastings and Mitchell, 1971; Reichelt ct ai, 1977; Jayabalan et ai, 1978) is held in numerous fine-diameter tubules that coalesce and empty into the esophagus via small ducts, thus allowing the bacteria to pass into the gut of the host (Harms, 1928; Haneda, 1940, 1950; Ahrens, 1965; Bassot, 1975). A facultative symbiont, P. leiognathi survives free of the host's light organ in various other marine habitats (Hastings, and Nealson, 1981). The behavioral value of luminescence, and the anatomical and physiological commitment of the fish to maintaining light organ bacteria, indicate the importance of bacterial light production to the fish. Consequently, given the fish's secondary control of the intensity and type of light emission by muscular shutters and chromatophores over the light organ, maximal luminescence would appear to be adaptive in this association. In this regard, alternative rapid-growth and nutrient-limitation models for the ponyfish symbiosis have been proposed that are consistent with high levels of luminescence (Nealson, 1979). However, essentially no information is available on the intensity of bacterial luminescence in situ or on the control of luminescence by the fish at the primary (physiological) level. The present study addresses questions of the physiological and morphological state of symbiotic bacteria in the ponyfish light organ. Measurements of bacterial light production and oxygen uptake, as physiological parameters of bacterial activity in the functioning symbiosis, are reported. The morphology, spatial distribution, and population density of light organ bacteria from several different ponyfish are described and interpreted in light of a newly discovered structural feature of this symbiosis, the light organ saccule. The results indicate that ponyfish bacteria grow slowly but produce consistently and exceptionally bright luminescence in the light organ, possibly in response to both anatomical and physiological controls exerted by the host fish. The results also provide a basis for in vitro studies of bacterial luminescence and growth that may lead to a better understanding of the physiology of this symbiosis. Materials and Methods Collection and handling offish The fish examined in this study were identified from the descriptions by Munro (1964), Kuhlmorgen-hille (1974), James (1975), Abe (1976), Rau and Rau (1980), and Dunlap and Mcfall-ngai (1984). Ponyfish captured live from Manila Bay, Luzon and Bais Bay, Negros Oriental, Philippine Islands (Gazza achlamys, G. minuta, Leiognathus bindus, L. brevirostris, L. equulus, L. fasciatus, L. splendens, and Sccutor insidiator), and at Misaki, Honshu, Japan (L. nuchal is), were maintained in laboratory aquaria near the collection sites as described by Mcfall-ngai and Dunlap (1983) or were transported live to laboratory facilities at the University of California, Los Angeles and maintained under similar conditions. For morphological examination of bacteria, light organs from freshly sacrificed specimens of these fish and from fresh specimens of L. elongatus and L. rivulatus from Suruga Bay and the Sagami Sea, Japan, were preserved in potassium-phosphate buffered 10% formalin, pH 70. Only healthy specimens of the fish, held less than one week (typically 1 to 2 days), were used for


16 POLYPODIACE.E OF THE PHILIPPINE ISLANDS. (1) D. lunulata Desv. Rhizome erect, rising above the ground or not; stipes clustered, fronds 1 to 2 m. high, commonly bipinnate; pinnules almost dimidiate, trapezoidal, the lower margin almost straight, the upper slightly rounded, entire, glabrous, herbaceous; sori 4 to G to a pinnule, large, partly immersed. Luzon, Cuming 142, Steere; Mount Apo, Devore and Hoover 345, common at about 4,500 feet. Pantropic. (3) POLYSTICHUM Roth. Fronds at least pinnate, usually tough or rigid, with toothed and spinulose margins and the basiscopic half the less developed; veins free, or in one species anastomosing sparsely and irregularly; sori round, usually dorsal; indusium normally peltate, exceptionally reniform. A reasonably natural genus, but difficult to define because of the many directions in which single species or groups vary; related to both Nephrodium and Aspidium. Mostly terrestrial. 1. Pinnae articulate to rachis (Cyclopeltis) (1) P. presumium 1. Pinnae not articulate, veins free Eupolystichum) 2. Simply pinnate (2) P. auriculatum 2. Lower pinnae pinnate, sori costal or medial. 3. Teeth aristate (3) P. aculeatum 3. Teeth blunt or mucronate (4) P. obtusus 2. Lower pinnse pinnate, sori submarginal (5) P. amabile 2. Lower pinnae at least bipinnate. 3. Teeth mucronate or awned. 4. Lower pinnae lanceolate-deltoid. 5. Rhizome creeping (6) P. aristatum 5. Rhizome erect (7) P. coniifolium 4. Pinnae linear (8) P. horizontale 3. Teeth awnless (9) P. varium 1. Pinnae not articulate, veins inclined to anastomose toward the margin (10) P. falcatum (1) P. preslianum (J. Sm) Moore. Rhizome creeping or oblique, short, densely fibrillose; stipes clustered, 3 to 10 cm. high, naked or nearly so; frond 20 to 40 cm. high, 8 to 10 cm. broad, simply pinnate; pinnae jointed to the rhizome, broadly lanceolate, obscurely crenate, almost horizontal, acute, cordate on the lower and truncate on the upper side at the base, coriaceous, glabrous; veins forked about three times, the lower branches falling short of the margin; sori in 1 to 3 rows on each side of the costa, the costal row the most constantly complete. Masbate and Catanduanes, Baranda ; Tayabas, Merrill 3351. Malaya. A species very near this, but with thinner foliage and the pinnae not articulate to the rachis, occurs in Luzon and Mindanao, and probably throughout the Archipelago; the two are decidedly too alike for me to separate them generically, by recognizing J. Smith's Cyclopeltis. (2) P. auriculatum (Sw) Presl, var nervosum (Fee) Christ. Stipes fasciculate, decumbent, 10 to 15 cm. long, scaly below or throughout; frond about 30 cm. long, 5 cm. broad; pinnae numerous, subsessile, crenate,


43 FIELDIANA ZOOLOGY Published by CHICAGO NATURAL HISTORY MUSEUM Volume 42 August 22, 1958 No. 4 Philippine Zoological Expedition 1946-1947 NOTES ON PHILIPPINE MALLOPHAGA, I. Species from Ciconii formes, Anseriformes, Falconiformes, Galliformes, Gruiformes and Charadriiformes K, C. Emerson AND Ronald A. Ward INTRODUCTION This is the first installment of a report on the Mallophaga collected from avian hosts by the Philippine Zoological Expedition, 1946-47. Since Mallophaga are obligatory external parasites, and highly host-specific, their distribution is dependent upon the distribution of the hosts. A list of the species of Mallophaga in the Philippines could be compiled by referring to a list of hosts known to inhabit, or migrate through, this region. Such a list has not been included in this paper because many records and descriptions have been based on material obtained from skins in museums or from live hosts in zoological gardens, and many still await confirmation. This study has arisen as a subsidiary enquiry to bird collecting, and it constitutes the first opportunity to confirm many host records. All records which were obviously mislabeled or mixed during collection have been omitted. Some series have not been given specific determinations because studies now in progress by other workers include several generic revisions. Library of Congress Catalog Card Number: 58-1335 ftaipiliwi No. 847 49 v O Of si NATURAL ' --' HISTORY SJRVE


EMERSON AND WARD: PHILIPPINE MALLOPHAGA 55 There appear to be no external characters by which this series can be separated from those normally found on Dendrocygna viduata (Linnaeus), the type host of this species. Family LAEMOBOTHRIONIDAE Laemobothrion mjobergi Eichler Laemobothrion mjobergi Eichler, 1944, Deutsche Ent. Zeitschr, 1943: 64, figs. 11, 12. Four males and four females from Haliastur Indus intermedius Blyth (Brahminy kite) were collected in the Liguasan Marsh, Cotabato Province, Mindanao. Laemobothrion sp. One male from Butastur indicus (Gmelin) (gray-faced buzzard) was collected at Bugasan, Cotabato Province, Mindanao. Suborder ISCHNOCERA Family PHILOPTERIDAE Craspedorrhynchus pachypus (Giebel) Docophorus pachypus Giebel, 1874, Insecta Epizoa, p. 71. One male from Haliastur indus intermedius Blyth (Brahminy kite) was collected at Manila, Luzon. Craspedorrhynchus sp. Two males and two females from Spilornis cheela holospilus (Vigors) (serpent eagle) were collected in Davao Province, Mindanao. Cuclotogaster acuminatus (Piaget) Lipeurus acuminatus Piaget, 1885, Les Pediculines, Suppl, p. 70, pi. 7, fig. 6. One female from Coturnix chinensis lineata (Scopoli) (painted quail) was collected at Naic, Cavite. Degeeriella sp. One male and two females from Haliastur indus intermedius Blyth (Brahminy kite) were collected in the Liguasan Marsh, Cotabato Province; six males and six females from the same host were collected at Manila, Luzon.


Department of Zoology 391 Copan; six crocodile skulls, including one very large one, collected in the Philippine Islands, from Mr. A. W. Exline, of San Jos, Mindoro; eight specimens from Dr. W. P. Kennedy, of Baghdad, Iraq; three Bahaman fresh-water turtles of recently described species, from the University of Miami, Miami, Florida; and two fine pink rattlesnakes (the rare species Crotalus lepidus lepidus), from Dr. and Mrs. Paul Rudnick, of the Mcdonald Observatory, Fort Davis, Texas. As in previous years, the Chicago Zoological Society, the Lincoln Park Zoo, and the General Biological Supply House, of Chicago, have turned over important material to the Museum. A large and important gift was that of 8,424 preserved fishes from the Zoology Department of the University of Chicago. These were collected by students and members of the faculty over a period of years and are of much value in studies of the local fauna of Illinois and neighboring states. Added to these were 248 specimens collected in Indiana and presented by Dr. Hurst Shoemaker of Stanford University, California. For exhibition, a number of game fishes were donated. Mr. Leon Mandel, of Chicago, gave a large "Mako" shark which he secured off the north coast of Cuba. Colonel Warren R. Roberts, of Chicago, contributed a white marlin, and Mr. Al Pflueger, of Miami, Florida, gave a specimen of Allison's tuna, a long-finned variety of the yellow-finned tuna. Mr. Michael Lerner, well-known sportsman of New York, presented an excellent mounted specimen of North Atlantic broadbill swordfish caught on rod and reel by Mrs. Lerner off Cape Breton, Nova Scotia — the first swordfish ever thus taken by a woman angler in Canadian waters. The John G. Shedd Aquarium, Chicago, continued its courtesy of past years by contributing a number of fish specimens. Excepting six specimens received in exchange, all the somewhat limited accessions of insects were obtained through various donors. The most noteworthy acquisition was a lot of 543 named beetles, mainly from the Austrian Tyrol. These were received as a gift from Dr. Wolfgang Amschler, of Zeiyarn bei Cronach, Bavaria, Germany. Mr. Gordon Grant, of Los Angeles, California, continued to show his interest in the Museum by donating 161 insects that he collected in his vicinity. Through the kindness of Dr. Orlando Park, Evanston, Illinois, forty-five species of named New Zealand moths were added to the collection. Of the 1,698 specimens of lower invertebrates accessioned, some 1,200 were shells presented by Mr. Clark W. Walter, of Chicago.


34 labials. Body moderate; a slight fold on the side and another on the posterior side of the hind limb; body and throat covered with very small granular scales, largest on the snout. Ventral scales imbricate. Male with 7 — 19 femoral pores on each side. Fig. 21. Hemidactylus garnoti D. B. Lower view of tail X SV-I-Fig. 22. Hemidactylus garnoti D. B, Fore limb X 5Tail depressed, flat below, with sharp denticulated lateral edge; the scales on the upper surface small, equal; those below larger, imbricate, with a median series of transversely dilated plates (fig. 2i). Limbs moderate; digits with a rudiment of web, dilated, inner well developed (fig, 22); 5 — 7 oblique lamellae under the inner digits, 10 to 12 under the fourth finger, 11 to 14 under the fourth toe. Brownish-grey above, uniform or with brown and whitish spots. Lower parts whitish. Length of head and body 60 mm; tail 80 mm. Habitat: Nias ; Sumatra (Tilatang Fort de Kock Matur Indragiri, Bingin Telok, Palembang, Agam); Java (Buitenzorg, Malang, Willis Mts, 5000 feet); Borneo (Kina Balu). — Malacca; Burma; Sikkim; Philippines; Solomon Islands; New Caledonia; Hawai; Tahiti. A common house-gecko in Sumatra. 6. Hemidactylus platyurus (Schneider). Stellio platyurus Schneider, Amph. Phys. II 1790, p. 30. Hemidactylus platyurus Boulenger, Cat. Liz. I 1885, p. 143 (s. syn). Cosymbotus platyurus Barbour, Mem. Mus. Comp. Zool. Harvard Coll. XLIV 1912, p. 81. Snout longer than the distance between the eye and the ear-opening, one time and a half the diameter of the orbit; forehead concave; ear-opening small, oval, oblique. Rostral four-sided,


36 Flores; Gr. Bastaard; Sumba; Savu; Kei Islands; Aru Islands; Halmahera; New Guinea (Sorong). — Malacca; Burma; Himalayas; Ceylon; South China; Indo-china; Philippines. Doubtful species. Hemidactylus tristis, described by Sauvage, Bull. Soc. Philom. (7) III 1878, p. 49, is said to be very much like Frenatus, differing only in the following points no tubercles on the base of the tail ; granules of the tail much smaller. Probably H. frenatiis D. B. Habitat: New Guinea. 5. Mimetozoon Boulenger. (BOULENGER, Pioc. Zool. Soc. London p. 767, 1896). Digits webbed for two-thirds, strongly dilated, with two rows of lamellae beneath; the distal phalanges clawed, slender and compressed, rising from within the extremity of the dilated part. Limbs, sides of head, body and tail with a large dermal expansion, covered above with imbricate tetragonal scales. Body covered with juxtaposed granular scales and tubercles. Pupil vertical. Males with femoral pores. Distribution. Penang; Borneo. A single species. I. Mimetozoon craspedotus (Mocq). Hemidactylus craspedotus Mocquard, Le Naturaliste XII 1890, p. 145; Nov. Arch. Mus. (3) II 1890, p. 126, pi. VII fig. 2; Bull. Soc. Philom. Paris (8) IX 189697, p. 20. Mimetozoon floweri Boulenger, Proc. Zool. Soc. London 1896, p. 767, pi. XXXVI. Snout narrow, pointed, longer than the distance between the eye and the ear-opening, one time and a half the diameter of the orbit; forehead concave; ear-opening small, oval, horizontal. Head-scales small, largest on the snout. Rostral rectangular, twice as broad as high, with median cleft ; nostril bordered by the rostral, the first labial and three nasals. Eleven to twelve upper and eight to ten lower labials ; mental very large, triangular; two pair of chin-shields, the inner larger and forming a suture. Body depressed ; from axilla to groin a membrane borders the flanks, another the neck from behind


u; or absent; one postocular; no temporals; five upper labials, third and fourth entering the eye; mental in contact with the anterior chin-shields ; latter in contact with three lower labials and about twice as large as the posterior. Tail pointed. Scales in 15 rows; ventrals 129146; anal entire; subcaudals 10 — 28. Brown or black above, iridescent; a yellow collar sometimes present and a yellow vertical spot above the angle of the Fig. 61. Pseudorhabdium longiceps (Cant). Nat. size. Side view of head. mouth. Lower surface dark brown, ventrals and outer scales with pale brown posterior border. Length of head and body 195 mm; tail 35 mm. Habitat: Nias; Sumatra (Deli, Langkat, Indragiri, Singkarah, Gunung Sahilan, Ajerbangis); Borneo (Sarawak, Kuching, Penrissen Road, Simanggang, Sebruang Valley, Pontianak); Celebes (Macassar). — Philippines; Malay Peninsula; Penang; Singapore. 35. Brachyorrhus Boie. (BOIE, Isis, p. 519, 1827). Head not distinct from neck; eye small; pupil vertically elliptic; nasal small, single or divided; internasals sometimes fused; loreal absent; praeocular present or absent. Maxillary teeth 20 — 22, equal; posterior mandibular teeth shortest. Body round, covered with smooth scales without pits, in 17 or 19 rows; ventrals rounded. Tail short; subcaudals in two rows. Distribution. Java; Timor; Moluccas; New Guinea. A single species.


42 sharp, finely denticulated lateral edge ; covered above with very small flat scales, below with a median series of large transversely dilated plates. Limbs depressed ; a fold of the skin bordering the hind limb posteriorly ; digits short, webbed at the base ; the lamellae angular, divided by a median groove (fig. 26). Greyish or reddish-brown above, uniform or variegated with darker brown, sometimes with round white spots on the back in regular longitudinal series. Lower parts dirty white. Length of head and body 98 mm ; tail 96 mm. Habitat: Nias ; Engano ; Sumatra (Medan, Singkarah Redjang, Gunung Sahilan); Banka; Borneo (Pontianak, Fig. 28. Gehyra mutilata c-o tt (Wiegm). Chin-shields and Gunung Kenepai, Smtang, Rivers Ho-throat X 2. wong, Upper Mahakkam and Baram, Kuching, Kidi district. Banting, Kina Balu, Sandakan) ; Celebes (Rurukan, Manado, Buol, Kema, Macassar); Java (Buitenzorg Surabaja) ; Flores ; Rotti ; Timor ; Sumba ; Ombaai ; Timor Laut; Kei Islands; Aru Islands; Ambon 1 ; Goram ; Halmahera; Ternate; Batjan; Waigeu ; New Guinea (lake Sentani, Lorentz river, Merauke Kaiserin-augusta river Astrolabe Bay, Berlinhafen, Seleo Island, Katow). — Malacca; Andamans; Nicobars ; Burma; Siam; Ceylon; Philippines; Mascarene Islands; Seychelles; Madagascar; Mauritius; Mexico; Cuba; d'entrecasteaux Archipelago; Bismarck Archipelago; Polynesia. House-gecko. 5. Gehyra baliola (A. Dum). Ilcmuhutylits baliolus A. Dumerilcat. Meth. Rept. 1851, p. 38; Arch. Mas. VIII 185556, p. 461, pi. XVII fig. 2. Gehyra baliola Boulenger, Cat. Liz. I 1885, p. 150Head more long than broad, snout conical, cheeks not swollen; forehead concave; ear-opening large, oval, oblique. Head granular, the granules enlarged on the snout. Rostral horseshoe-shaped; nostril bordered by the rostral, the first labial, a


1 89 Sumba; Rotti; Timor; Wetar; Celebes (Manado, Kema, Kandari, Pare Pare, Raha); Sangir Islands; Ternate; Halmahera; Batjan; Buru; Ceram; Ambon; Saparua; Coram; New Guinea. — Pelew Islands; Malay Peninsula; Penang; Andamans; Nicobars; Siam; Indo China; Burma; Ceylon; India; Philippines; N. Australia. Not rare at the sea-coast; feeds on fish. 41. Fordonia Gray. (Gray, Zool. Misc. p. 67, 1842). Head small, not distinct from neck; eye very small; pupil vertically elliptic; shields of head large; nasals undivided or partly divided, separated by a single internasal; no loreal. Maxillary teeth small, 7 or 8, followed by two enlarged, grooved teeth; mandibular teeth about equal. Body round, covered with smooth scales, without pits, in 25 — 29 row; ventrals rounded. Tail short, subcaudals all or partly in two rows. Distribution. Burma; Cochin China; Indian Archipelago; N. Australia. A single species. I. Fordonia leucobalia (Schleg). Homalopsis leucobalia Schlegel, Phys. Serp. II 1837; p. 345, pi. XIII, fig. 8, 9. Fordonia leucobalia Boulenger, Cat. Sn. Ill 1896, p. 21 (s. syn). Fig. 71. Fonioiiii leucobalia (Schleg) X V41 view of head.


INGER: AMPHIBIA, PHILIPPINE EXPEDITION 223 Fig. 37. A, Megophrys monticola stejnegeri ( X 06) ; B, M. m. nasuta ( X 04) ; C, M. m. monticola (X 05); D, M. m. ligayae (X 05). one pair of longitudinal dermal folds on back; folds reaching end of trunk (fig. 37). Range. — Java. Megophrys monticola nasuta Schlegel Ceratophryne nasuta Schlegel, 1858, Handl. Dierk, 2: 57, pi. 4, fig. 72 — Sumatra. Megophrys nasuta Smith, 1930, Bull. Raffles Mus, no. 3, p. 132; 1931, op. cit, no. 5, p. 9; Mertens, 1934, Arch. Hydrobiol, 12, Suppl. (Trop. Binnengew), 4680. Megalophrys nasuta Boulenger, 1908, Proc. Zool. Soc. London, 1908: 413, pi. 22; van Kampen, 1923, Amph. Indo-austr. Arch, p. 10, fig. 2. Megalophrys montana Boulenger, 1908, Proc. Zool. Soc. London, 1908: 411 (part); van Kampen, 1923, Amph. Indo-austr. Arch, p. 8 (part). Megophrys monticola Smith, 1931, Bull. Raffles Mus, no. 5, p. 12; Mertens, 1934, Arch. Hydrobiol, 12, Suppl. (Trop. Binnengew), 4: 679 (part). Material examined. — Borneo, 13 (10 MCZ; 3 UMMZ). Diagnosis. — A large form of monticola; adult females about 110 mm, males 90 mm. (see Table 3) ; dermal appendage of snout usually well developed (up to 35 mm), occasionally present only as a small projection; orbital appendage large (35 to 70 mm); two pairs of


INGER: AMPHIBIA, PHILIPPINE EXPEDITION 229 to 75 mm, snout to vent). The parietal crests of specimens from the Calamian Islands are on the average thicker than those on material from Palawan and Balabac. This difference is correlated with differences in body lengths and may not indicate a change in growth relations. Another characteristic of the supraorbital-parietal crest showing geographic variation is the position of its highest point. In the specimens from Java, Bali, and Borneo the high point is opposite the posterior third of the eye; in all specimens from the Philippines the high point is behind the eye, that is, on the parietal portion of the crest. The parotoid glands of the two subspecies are different in shape. Specimens of biporcatus from Java, Borneo, and Bali have subtriangular parotoids, the hind end being decidedly narrower than the front end (fig. 38, B). Occasionally the parotoid in these is attentable 6. Geographic variation of characters in adult Bufo biporcatus


MARINE ANIMALS THAT ARE POISONOUS TO EAT 125 Snapper, Monotaxis grandoculis (Forskal) (Fig. 82, Top) Polynesia, westward to east Africa. Chinaman Fish, Paradicichthys venenatus Whitley (Fig. 82, Center). Australia. Parrotfish, Scars caeruleus (Bloch) (Fig. 82, Bottom). Florida and West Indies. Parrotfish, Scams microrhinos Bleeker (Fig. 83, Top). Indo-Pacific. Seabass, Grouper, Cephalopholis argus Bloch and Schneider (Fig. 83, Next to Top). Tropical Indo-pacific. Seabass, Plectropomus oligacanthus Bleeker (Fig. 83, Next to Bottom). Indonesia, Philippine, Caroline and Marshall Islands. Seabass, Plectropomus truncatus (Fowler) (Fig. 83, Bottom). Micronesia, Indonesia, Philippines. Seabass) Variola loutiforska) (Fig. 84, Top). Tropical Indo-Pacific. Barracuda, Sphyraena barracuda (Walbaum) (Fig. 84, Bottom). Indo-pacific; from Hawaii to the Red Sea; west Atlantic from Brazil to the West Indies, Florida and Bermuda. Medical Aspects. Tingling about the lips, tongue, and throat, followed by numbness may develop immediately or any time within a period of 30 hours after ingestion of the fish. The tingling sensation may be accompanied by such other symptoms as nausea, vomiting, metallic taste, dryness of the mouth, abdominal cramps, and diarrhoea. The muscles of the mouth, cheeks and jaws may become drawn and spastic, with a feeling of numbness. Headache, joint aches, nervousness, prostration, dizziness, pallor, cyanosis, inability to sleep, extreme weakness, and exhaustion are frequently present. The feeling of weakness may become progressively worse until the patient is unable to walk. Muscle pains are generally described as dull, heavy aches or cramping sensations, but may also be sharp, shooting, affecting particularly the arms and legs. Victims complain of their teeth feeling loose and painful in their sockets. Visual disturbances consisting of blurring, temporary blindness, sensitivity to light, and spots before the eyes are not uncommon. Skin disorders are frequently reported consisting of intense itching, red popular rash, blisters, extensive areas of loss of skin, especially of the hands and feet, and occasionally ulceration. There may also be loss of hair and nails. In severe intoxication the nervous symptoms are particularly pronounced. The victim may interpret the feeling of cold as a Fig. 79. Top to bottom: Squirrelfish, Myripristis murdjan (Forskal). Oceanic Bonito, Katsuwonus pelamis (Linnaeus). Wrasse, Epibulus insidiat or (Pallas). (From Hiyama) Wrasse, Julie gaimardi (Quoy and Gaimard). (Arita)


MARINE ANIMALS THAT ARE POISONOUS TO EAT 133 POISONOUS MARINE TURTLES Poisoning from marine turtles is one of the lesser known types of intoxications produced by marine organisms. The cases that have been reported are sufficiently severe to be impressive. As in the case of fishes, most of these species are commonly eaten with impunity. For some unknown reason, certain species of marine turtles in the vicinity of the Philippine Islands, Ceylon, and Indonesia, under certain circumstances, may become extremely poisonous to eat. Species of Marine Turtles Reported as Poisonous to Eat Green Sea Turtle, Chelonia my das (Linnaeus) (Fig. 85, Top). Inhabits all tropical and subtropical seas. Hawksbill Turtle, Eretmochelys imbricata (Linnaeus) (Fig. 85, Center). Inhabits all tropical and subtropical seas. Leatherback Turtle, Dermochelys coriacea (Linnaeus) (Fig. 85, Bottom). Largely circumtropical, but occasionally taken in temperate waters. Medical Aspects. Symptoms generally develop within a few hours to several days after ingestion of the flesh. The initial symptoms are usually nausea, vomiting, diarrhea, severe upper abdominal pain, dizziness, dry burning sensation of the lips, tongue, lining of the mouth and throat. Swallowing becomes very difficult, and excessive salivation is pronounced. The disturbances of the mouth may take several days to develop, but become progressively severe as time goes on. The tongue develops a white coating, and the breath becomes very foul. Later, the tongue may become covered with multiple pinhead-sized, reddened papules, which may later break down into ulcers. If the victim has been severely poisoned, he tends to become very sleepy, and is difficult to keep awake. If this symptom develops, it is usually a bad sign, and death soon follows. Death is believed to be due to liver and kidney damage. About 44 percent of the victims poisoned by marine turtles die. Treatment. There is no specific treatment. Some of the recommendations presented in fish poisoning are pertinent here. Prevention. Marine turtles in the tropical Indo-pacific region should be eaten with caution. If in doubt, check with local native groups and find out if they are safe to eat in that locality. Turtle liver is especially dangerous to eat. Fig. 85. Top: Green Sea Turtle, Chelonia mydas (Linnaeus). Center: Hawksbill Turtle, Eretmochelys imbricata (Linnaeus). Bottom: Leatherback Turtle, Dermochelys coriacea (Linnaeus).