13 which was taken on Ticao April 17, 1902. The species should be added to the list of birds from that island. ^GIALITIS DUBIA (Scop). Fairly abundant on Calayan. aegialitis alexandrina (Linn). Fairly abundant on Calayan. AEGIALITIS PERONI (Bp). Fairly abundant along beaches on both Fuga and Calayan. OCHTHODROMUS MONGOLUS (Pall). Two specimens from Calayan. Heteractitis brevipes (Vieill). One killed on Calayan October 21 was the only specimen seen. A female was collected on Ticao May 27, 1902, and is to be added to the Ticao list. Tringoides hypoleucus (Linn). Abundant on Calayan. Rhyacophilus glareola (Gm). One specimen from Calayan. LIMONITES RUFICOLLIS ( Pall. ) Very rare; one killed October 23 on Calayan. GALLINAGO STENURA (Kuhl). Snipe were abundant on Calayan, and the only one preserved is of this species, of which the only previous Philippine record is Mindanao. About Manila Gallinago megala Swinh. is the common species. Two specimens of G. gallinago (Linn) were taken by us at Pandacan, a suburb of Manila, but stermra appears to be rare. Key to the Philippine species of Gallinago. A. Tail feathers, 20 or more, the outer ones stiffened and very narrow; dark bars on axillaries wider than the white bars. a. Tail shorter, about 190 inches; outer feathers narrower G. genera. 6. Tail longer, about 210 inches; outer feathers wider G. megala. B. Tail feathers, usually 14, normal; dark bars of axillaries narrow or obsolete '' O-lhrwgo. Nycticorax nycticorax (Linn). A dozen or more roosted during the day in a thicket of trees near the beach. A young bird was preserved. Another was killed as it sat alone in some brush near a small stream. Calayan only. MESOPHOYX INTERMEDIA Wagl. One or two pairs on Calayan feeding in same fields as Jmmlcus. One female collected.


14 Demiegretta sacra (Gm). Observed about the beaches of Fuga and Calayan. BUBULCUS COROMANDUS (Bodd). A small band always to be seen in company with the cattle on Calayan. Local name, "la-guava' The species was not noted on Fuga. BUTORIDES JAVANICA (Horsf). Ear; one specimen from Calayan. Ardetta sinensis (Gm). From Calayan only, where a live bird was brought to us October 23. Local name, "rat' POLIONETTA ZONORIIYNCHA (Swinh). Anas zonorhyncha Salvadori, Cat. Bds, XXVII, p. 211. On December 7 a dozen birds of this species rested on a small river near the beach, and a single female was killed. This is probably its most southern capture. Bill jet black with a broad tip of bright yellow ; most of the nail black; irides tan brown; legs and feet light salmon; nails black; webs dusky. QUERQUEDULA QUERQUEDULA (Limi). A male in ragged plumage was collected October 29 on Calayan. Nettium crecca (Linn). Querquedula crecca Whitehead, Ibis, 1899, p. 501 (Benguet Province). Secretary Worcester killed two males in Bulacan, Luzon, in January, 1903. A male, October 2G, and a female, November 10, were killed on Calayan. FULIGULA FULIGULA (Linn). Salvadori says of this species: Accidental in the Malay Archipelago (Philippines and Borneo)" I believe, however, that it is a regular fall and winter visitant to our northern islands at least. We have specimens from the Laguna de Bay, Luzon, and it was common on Calayan considering the scant accommodation for ducks. I have seen numbers of them in Quinta Market, Manila. Parties of from two to five could usually be found on a small river or some of the carabao wallows, where they feed on small freshwater mollusks or the larvae of frogs. Mareca PENELOPE (Linn) ; Salvadori, Cat. Bds, XXVII, p. 231. A male from Calayan, November 2G. No more were seen. Previously unrecorded from the Philippines. Spatula clypeata (Linn). One specimen from Calayan. "Dulumpapa" is the name used for any species of the duck tribe. PELECANUS PIIILIPPEN8TS Gm. The Philippine pelican has been recorded but rarely from the Islands and probably it is strictly confined to fresh-water marshes and lakes


87 In Figure 4a, a decrease in coral cover in 2005 is evident only at Site 5 but this cannot be attributed to the tsunami and is most likely related to a mild bleaching event in 2003. At site 3, coral cover is as high in 2005 as has ever been recorded at this location since 1989. Figure 4b shows no loss of cover as a result of the tsunami at sites 9 and 1 1 although there was a marked drop in coral cover at site 9 between 1990 and 1991 as a result of an extensive bleaching event in 1991. There has been very little recovery at this location in subsequent years. At intertidal site 10, coral cover was lowest in 1997-98 during a period of exceptionally low sea level. Field observations at this location revealed no physical damage as a result of the tsunami and this was reflected in the high coral-cover values of 2005. Generally diversity indices showed very little change over time at both affected and unaffected sites (data not shown) with no clear patterns emerging at sites affected by the tsunami. DISCUSSION The Indian Ocean tsunami of 2004 clearly had a limited effect upon the coral reefs of the Andaman Sea coast of Thailand. Remarkably, there appears to be few references to the effects of tsunamis on coral reefs in the literature despite a history of repeated tsunamis in the Indo-pacific region. For example, a total of 35 tsunamis have been estimated to have impacted the Indonesian archipelago since the Krakatau tsunami of 1833 (Carey et al, 2001) while significant tsunami waves were reported following earthquakes at Car Nicobar in 1881 and in the Andamans in 1941 (Bilham et al, 2005). Coral reefs were mentioned in a report of a tsunami initiated as a result of an earthquake in the Philippine Fault Zone in S.E Mindanao in 1992 but only in terms of their ameliorating effects in reducing the wave height finally reaching the shore (Besana et al, 2004). Although the heights of the tsunami waves are not reflected in the tidal measurements obtained for the relevant period at Ko Taphao Noi, Harada (2005) estimates tsunami wave heights to have been approximately 1 0m on the mainland inshore from sites 7 and 8, 3 m at sites 9, 10 and 1 1 and 5 m at sites 15 and 16. These heights were measured on site within four days of the arrival of the tsunami waves. Coral reef damage appears to have been mainly restricted to sites on the west-to-southwest sides of islands which are frequently exposed to southwest monsoon influences. Coastal topography and aspect of site similarly played an important role in influencing tsunami-related damage to coral reefs in northern Sumatra in December 2004 (Baird et al. 2005). While poorly attached massive corals at depth were displaced in Sumatra (Baird et al. 2005) damage was mainly restricted to shallow reef sites in Thailand. At the few locations where negative impacts were observed along the Thai coastline, the type of damage noted was similar to that of hurricanes and cyclones with broken branching corals (Woodley et al, 1981, Woodley, 1993; Rogers. L993) and dislodgement of often weakly attached massive colonies (Massel and Done. 1993) in shallow waters. Similar dislodgement of large colonies of Acropora palifera has been


LIST OP PHILIPPINE BIRDS. -i and lucidus, having the back of one and the ventral surface of the other, but the crimson marking of the face and throat separates it from either. 52. Tiga Everettii. Palawan. 53. Yungipicus maculatus. Panay. 54. Yungipicus validirostris. Mariuduque, Luzon. It seems most probable that it was the Luzon bird described by Blyth under this name. 55. Yungipicus Basilanicus. sp. nov. Basilan. Head black slightly tinged in front with cinnamon brown. Crimson crest In male nearly confluent. Breast washed with orange red, fading to sulphur yellow behind. Interscapulars black and barred with sulphur yellow. Back sulphur yellow. 56. Yunigipicus Leytensis. sp. nov. Head black. Forehead and basal region cinnamon. Breast tinged with crimson. TROGONIDJE. 57. Harpactes ardens. Mindanao, Basilan, Samar, Marinduque, Luzon. MEROPIDAE. 58. Merops Philippinus. Mindanao, Basilan, Guimaras, Panay, Negros, Cebu, Siquijor, Panay, Luzon. 59. Merops bicolor. Mindanao, Guimaras, Negros, Marinduque, Mindoro, Luzon. Nesting in Mariuduque in May in open, nearly level ground. Merops Philippinus feeds solitarily upon wasps, dragon flies, etc, M. bicolor in flocks and apparently in nearly all cases upon honey bees. CORACTDIE. 60. Eurystomus orientalis. Mindanao, Basilan, Guimaras, Panay, Negros, Siquijor, Samar, Leyte, Mariuduque, Mindoro, Luzon. ALCEDINID.E. 61. Alcedo Bengalensis. Paragua, Balabac, Mindanao, Basilan, Guimaras, Negros, Siquijor, Samar, Masbate, Luzon. 62. Pelargopsis gigantea. Mindanao, Basilan, Samar, Guimaras, Masbate. 63. Pelargopsis leucocephala. Paragua, Mindoro. 64. Ceyx melanura. Luzon. The Melanura type of Ceyx takes on a number of representative


6 LIST OP PHILIPPINE BIRDS. PSITTACIDAE. 2. Prioniturus discurus. Mindanao, Basilan, Guimaras, Leyte, Samar, Masbate. 3. Prioniturus cyaneiceps. Paragua. 4. Prioniturus Luconensis. sp. nov. Marinduque, Luzon. Adult male and female. General color above and below, including the head, bright yellow green. 5. Prioniturus Mindorensis. sp. nov. Mindoro. Adult male. General color, grass green as in P. discurus ; crown and nape, violet; lores, face and chin, yellow green. 6. Tanygnathus Luconensis. Paragua, Basilan, Guimaras, Panay, Negros, Masbate, Marinduque, Mindoro. 7. Tanygnathus Everettii. Mindanao. 8. Cyclopsitta lunulata. Luzon. 9. Cyclopsitta Mindanensis. sp. nov. Mindanao. Adult male. Collar violet, cheeks green. C. lunulata has a cobalt collar and blue cheeks in the adult male. 10. Loriculus Philippensis. Marinduque. 11. Loriculus regulus. Panay, Guimaras, Negros, Masbate. L. Panayensis of Tweedale is the young male of L. regulus. 12. Loriculus Hartlaubii. Mindanao, Basilan. 13. Loriculus chrysonotus. Cebu. 14. Loriculus Siquijorensis. sp. nov. Siquijor. Adult male with large quadrangular spot of crimson upon the forehead and crown, ending sharply in the general green color of the upper surface. Adult female and young male: Red spot upon the forehead veiled by blue tips to the red feathers. 15. Loriculus Mindorensis. sp. nov. Mindoro. Adult with a broad crimson band across the forehead below the eyes, ending sharply in the green of the upper surface. 16. Loriculus Worcesteri. sp. nov. Samar, Leyte. The crimson of the upper surface covering the forehead, crown and nape, where it tapers to a point. It is distinguished from L. Hartlaubii, its nearest ally, by the much fainter wash of orange red upon the mantle and interscapulars, and by the shape of the red markings of the head and nape. In L. Hartlaubii the crimson area grows wider on the nape, where it ends broadly. All adult male Philippine Loriculi observed, have an orange-red spot upon the breast, and green cheeks. All adult females and all young males in their second plumage have blue on the cheeks and throat, and lack the orange-red plastron. All young birds in first plumage have the entire head green, and the rump scarlet.


4 INTRODUCTION. esis of Mr. A. E. Wallace, founded on the former meager knowledge of the Philippine fauna, that the islands were oceanic, and never had a connection with the continent. The relationship of the tamarou with the area of Celebes, and the general similarity of the Philippine and Celebes birds and mammals, especially the latter, would seem to point to a former connection with that island, and the ancient existence of a Celebes-Philippine continent lying outside of Java and Borneo. Over fifty supposed new species of birds and mammals were discovered by the expedition, and are here briefly described. This work is necessarily more or less imperfect and faulty from the lack of the complete literature of the subject, as well as from the lack of large collections of Philippine birds in this country with which to compare those in our hands. It is hoped, however, that the results of our labor will be found to make a real and valuable addition to the knowledge of this interesting group of islands.


INTRODUCTION. The Steere Expedition to the Philippines went out from the University of Michigan in the year 1887, and spent about twelve months in the Islands. The object of the expedition was to make general zoological collections, at the same time visiting as many distinct localities as possible, so that the distribution of species in the islands themselves could be studied. Fifteen of the larger islands, situated in all parts of the group, were visited, and from two to six weeks spent upon each. This amount of time, with a party of five collectors from the United States, and such native help as could be obtained, sufficed to make very large, though by no means exhaustive collections of vertebrates, and important collections in several groups of invertebrates. As island after island was reached and collected upon by the party, the discovery was made that the group is divided into several quite distinct zoological subdivisions, which I have considered to be sub-provinces, and have named as follows 1st, the sub-province of North Philippines, made up of Luzon, and small adjacent islands ; 2d, Mindoro ; 3d, Central Philippines, embracing the islands of Panay, Negros, Guimaras, Cebu, and Masbate ; 4th, West Philippines, Paragua and Balabac ; 5th, South Philippines, Mindanao and Basilan; 6th, East Philippines, Samar, Leyte, and probably Bojol. There are marks of still further differentiation within these divisions, notably in the instances of Cebu and Basilan. The discovery by the expedition in the centrally placed island of Mindoro of a new species of water buffalo, the tamarou, an animal of such habits and of such large size as to make its introduction by man improbable, goes far toward destroying the hypoth


THALAMIFLORiE. 1 1 50. Tamarix orientalis, L. "Khagal or Tarash" a. Leafy twig. Stewart, ' Punjab Plants,' p. 92. HYPERICINEAE. 51. Vismia dealbata, H.B.K. " Lacre-zeira." a. Flowering branchlet. b. Twig with young fruit. Santarem, Para; H. W. Bates, Aug. 18, 1853. The following note is attached to the specimen a. : — « "No. 8, Lacrezeira, with specimens of gum and roots. The specimen of gum I collected from one wounded tree. The tree « as slender ! height, 20 feet ; diameter of stem, 4 inches. It runs slowly like wax, but soon solidifies. It is an indelible pigment, colour of gamboge. I supposed the tree was a species of Vismia. The roots, I now find, are much esteemed as a remedy here ; decoctions drank in tea have performed great cures in old chronic gonorrhoea. The botanical specimens were taken, some from the tree, which yielded the gum, and others from those from which I took the roots ; not observing any specific difference, I have put them together." Specimen b is labelled, " Yields a red gum, applied externally in ringworm, &c. Received March, 18.53." GUTTIFEK.*:. [52. Garcinia Hanburyi, Hook, f.] Garcinia Morella, Desrouss. var. pedicellata. a. Leaves. Singapore, D'Almeida. With 6 drawings attached, including a sketch of the tree, leaves, flowers, and dissections of the flower and fruit. [These drawings formed the basis of the illustrations published in the Trans. Linn. Soc, 1864, vol. xxiv. tab. •50a, p. 487. The plant was subsequently raised to the rank of a species by Sir J. D. Hooker, tuider the name of G Hanburyi, Hook./. See Journ. Lin. Soc, vol. xiv., p. 48.5, 1875. Bentl. and Trim. Med. PL, tab. 33. Hanbury 'Science Papers,' p. 326-333.] DIPTEROCARPE^. 53. Dryobalanops Camphora, Colebr. A leafy twig bearing flower buds and detached leaves and flowers. Sumatra, Dr. W. H. K. De Vriese, Leyden, 8 Aug., 1857.


32 PHILIPPINE BAMBOOS Ranger Mabesa has made counts of the number of shoots produced in one season on two areas of Schizostachyum lumampao, each area consisting of 100 square meters. The two areas together contained 340 canes and produced 35 shoots, or 10 per cent of the original number. All canes were cut from two smaller-sized plots in May, 1917, and the number of new canes counted in November, 1917. These two plots originally contained 221 canes. After the clearing they sent up a large number of shoots about the size of a lead pencil and 10 usable canes, which is only 45 per cent of the original number. This would indicate that it requires some years for this bamboo to regain its original density when once completely cut-over. COST OF HARVESTING AND PRICES Data on the cost of harvesting were collected during the year 1916 on the College of Agriculture farm at Los Banos, Laguna. One man was employed for a total of 91 days, during which time 1,000 canes of Bambusa spinosa were harvested, an average of 11 canes per day. The cost of labor was 80 centavos per day, making the total cost of harvesting 1,000 canes 7280 pesos or an average per cane of 728 centavos. In the same locality these canes sold at 4 pesos per dozen. Bamboo sold in Manila is floated down the river from the country behind or across Manila Bay. In Manila the present prices for canes in the river are: First-class canes, 32 pesos per hundred; second-class canes, 27 pesos per hundred; and third-class canes, from 18 to 20 pesos per hundred. The cost of floating bamboo to Manila is very small, as rafts containing large numbers of canes can be operated by a very few men.


PLATE XX. GUADUA PHILIPPINENSIS GAMBLE. COTYPE SPECIMEN.


SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES coral and the coralline algae species tested. The specificity of bacterial communities to different coralline algal species has rarely been investigated (Johnson et al, 1991). With the recent development of more refined genetic techniques it is possible to compare different microbial communities, which might enable the identification of the microbes) that can induce coral larval settlement and metamorphosis. Negative Settlement Cues Water quality and substrate conditions impact fertilization rates and also may inhibit some coral larvae from normal settlement and metamorphosis. Low coral recruitment is commonly documented in the field, yet surprisingly few studies have experimentally tested which substrate characteristics might deter coral larval settlement. Coral larval survival and settlement can be reduced by many environmental stresses, such as elevated temperatures (Edmunds et al, 2001), variation in salinity (Vermeij et al, 2006), sedimentation (Hodgson, 1990; Gilmour, 1999), and UVB radiation (Kuffner, 2001; Gleason et al, 2006). Survival and settlement are reasonable ecological metrics for the effects of stress, but an important gap in our knowledge is how sublethal stress influences larval behavior and post-settlement health and success (Downs et al, 2005). New techniques including cellular biomarkers and differential gene expression using microarrays should provide important techniques to measure sublethal stress in coral larvae. Water quality conditions that are known to impact adult corals also have dramatic effects on larval supply and settlement. Of the physical conditions that negatively influence larval settlement, elevated temperature has received the most attention and has the potential to increase in frequency and duration as ocean temperatures continue to warm. Larvae of the Caribbean brooding coral Pontes astreoides were killed and had low densities of zooxanthellae when exposed to elevated temperatures for 24 hours (Edmunds et al, 2001, 2005). High temperatures (36C) killed Acropora muricata larvae within 40 hours (Baird et al, 2006), and temperatures of 32C killed Diploria strigosa larvae and reduced their settlement (Bassim and Sammarco, 2003). However, at elevated temperatures (29C) larvae of Stylophora pistillata had the same settlement as at 25C (Putman et al, 2008), and more larvae settled on the CCA in 25C than in 23C. Many of these studies used different experimental conditions, making it difficult to compare the effects of temperature on different species of coral larvae. Temperature is one stress that is relatively well studied, but more research is necessary to understand other physical stresses, such as ocean acidification (Albright et al, 2008), that will affect coral larvae in the future. Larval interactions with the biological inhabitants of reef communities can also reduce larval settlement. Algal turfs, macroalgae, and benthic cyanobacteria can negatively impact the settlement of coral larvae (Kuffner and Paul, 2004; Birrell et al, 2005; Kuffner et al, 2006; Birrell et al, 2008a). In the Florida Keys, two brown algae, Dictyota pulchella and Lobophora variegata, reduced the total number of Pontes astreoides settlers (Kuffner et al, 2006). In the Philippines, the algae Sargassum polycystum and Laurencia papulosa decreased larval settlement of Pocillopora damicornis, but water conditioned with these algae increased settlement over the seawater controls (Maypa and Raymundo, 2004). In Australia, water conditioned with the foliose brown alga Padina sp. reduced larval settlement of Acropora millepora; however, water conditioned with the brown alga Lobophora variegata increased settlement (Birrell et al, 2008a). The cyanobacterium Lyngbya majuscula reduced the survivorship of Acropora surculosa larvae and settlement and metamorphosis of Pocillopora damicornis in studies conducted on Guam (Kuffner and Paul, 2004), and in Florida, the cyanobacterium Lyngbya polychroa caused Pontes astreoides to avoid settling adjacent to it on settlement tiles (Kuffner et al, 2006). Some macroalgae and cyanobacteria can act as settlement inhibitors for coral larvae, but this was not true for all the algae tested. A surprising contrast was observed for Favia fragum larvae, which had high rates of settlement and metamorphosis onto live Halimeda opuntia when offered with coral rubble (Nugues and Szmant, 2006). Coral larvae of Montipora capitata were observed to settle onto Viva sp. (Figure 4e; Vermeij et al, 2009). Why these larvae would settle directly onto blades of algae is unclear as this substrate is ephemeral, thus probably increasing post-settlement mortality. Little research has been done on the mechanisms that algae use to inhibit settlement, but algal qualities such as natural products, shading and abrasion, serving as vectors of bacteria, and releasing dissolved organic matter may contribute to the negative impacts of algae on larval settlement. Competition from other members of coral reef communities also influences larval behavior. Tissue of the scleractinian coral Goniopora tenuidens suspended in seawater inhibited metamorphosis of Pocillopora damicornis larvae and reduced the growth of new recruits over seven days (Fearon and Cameron, 1996). The tissue from Goniopora tenuidens also caused increased mortality of larvae from P. damicornis, Platygyra daedalea, Fungia fungites, and Oxypora lacera. Increased research on the types of benthic


Page 404 The Veliger, Vol. 44, No. 4 San Luis Obispo County, and eradication efforts undertaken as necessary. The potential is great for harm by R. clecollata to the unique, endemic land snail fauna of San Nicolas Island. Among the strictly endemic native snails, Micrarionta opuntia Roth 1975, never attains a shell diameter of more than about 103 mm; the mean adult shell diameter of Micrarionta feralis (Hemphill, 1901) is around 15 mm (Pearce, 1990; Roth, 1996). For these species, the "refuge" of a size greater than 15 mm diameter (which allows Helix aspersa to persist at low numbers in the presence of R. decouata) is unavailable. In addition, the total known range of M. feralis covers only a few hundred square meters. The San Nicolas Island species Xerorionta tryoni (Newcomb, 1864) attains an adult shell diameter of 1928 mm. Size-frequency distribution of a population observed in 1981 suggests that maximum shell size and sexual maturity are attained late in the second or early in the third year of life. A shell diameter of 15 mm is probably not attained until the second year of life (BR, unpublished observations). Time to maturity in Helix aspersa varies, but in a coastal southern California setting with artificially applied water, snails matured in 6-8 months (Potts, 1975). Under laboratory conditions, a shell diameter of 15 mm was attained in about 25-45 days (Potts, 1975: fig. 2). Juvenile Xerarioiita tryoni spend more time in the vulnerable 15-mm size class than do juvenile Helix aspersa, and therefore the potential impact of predation upon populations of X tions is greater than that recorded for H. aspersa. The apparent localization and slow spread of Riimino decollata on San Nicolas Island suggest that its eradication on that island may be possible. Acknowledgments. Jim Carlton. Rob Cowie. Tim Pearce, and David Robinson read and commented on a draft of the manuscript. Jim Strampe helped produce the California distribution map, and Frank Focha and John Warrick (CDFA) helped interpret the pertinent regulations. We are grateful to Tim Pearce, Sally Walker, Ken Warheit, Alicia Cordero, David Lindberg, and others who helped search for R. decollata on San Nicolas Island. Literature Cited Batts, J. H. 1957. Anatomy and life cycle of the snail Riiiuina decollata (Pulmonata: Achatinidae). Southwestern Naturalist 22-3)74-82. California Department of Food and Agriculture (CDFA). 1998. Plant Quarantine Manual. Pest Exclusion Branch, Plant Health And Pest Prevention Services Division. Animal Policy section updated on 12 January 2000. Dundee, D. S. 1970. Introduced Gulf Coast molluscs. Tulane Studies in Zoology and Botany 173): 101-1 15. Dundee, D. S. 1974. Catalog of introduced mouusks of eastern North America (north of Mexico). Sterkiana 551-37. FISHER, T. W. 1966. Rumina decollata (Linnaeus, 1758) (Achatinidae) discovered in southern California. The Veliger 91): 16. Fisher, T. W. & R. E. Orth. 1985. Biological control of snails. Occasional Papers, Department of Entomology, University of California, Riverside 1. viii H1 11 pp. Fisher, T. W, R. E. Orth & S. C. Swanson. 1980. Snail against snail. California Agriculture 34ll-12)18-20. Pearce, T. A. 1990. Phylogenetic relationships of Micrarionta (Gastropoda: Pulmonata) and distinctness of the species on San Nicolas Island. California. Malacological Review 23137. Pearce, T. A. 1993. Micrarionta (Gastropoda: Pulmonata on San Nicolas Island, California. Evolutionary relationship among the species. Malacological Review 2615-50. Potts, D. C. 1975. Persistence and extinction of local populations of the garden snail Helix aspersa in unfavorable environments. Oecologia 21313-334. Robinson, D. 1999. Alien invasions: the effects of the global economy on non-marine gastropod introductions into the United States. Malacologia 41413-438. Roth, B. 1996. The fossil land snail Micrarionta (Micrarionta) intermedia Pilsbry (Gastropoda: Pulmonata): insular phylogeny revisited. The Veliger 393)204-212. Roth, B. & C. M. Hertz. 1997. Recent records of Cochlicella barbara (Linnaeus, 1758) (Hygromiidae) in southern and central California. The Festivus 299)81-83. Selander, R. K. & D. W. Kaufman. 1973. Self-fertilization and genetic population structure in a colonizing land snail. Proceedings of the National Academy Sciences of the U.S.A. 704): 1186-1 190. Selander, R. K. & R. O. Hudson. 1976. Animal population structure under close inbreeding: the land snail Rumina in southern France. American Naturalist 1 10974)695-718. Selander, R. K, Kaufman, D. W. & R. S. Ralin. 1974. Self-fertilization in the terrestrial snail Rumina decollata. The Veliger 163)265-270. The Occurrence of the Shell-less Neritacean Gastropod Titiscania limacina in the Galapagos Islands Jose Templado Museo Nacional de Ciencias Naturales (CSIC), Jose Gutierrez Abascal, 2, 28006 Madrid, Spain and Jesiis Ortea Departamento de Biologia de Organisms y Sistemas, Laboratorio de Zoologia, Universidad de Oviedo, 33071 Oviedo, Spain Only two species of shell-less neritacean gastropods of the family Titiscanidae have been described: Titiscania limacina Bergh, 1875, and T. shinkishihataii Taki, 1955. The former was discovered and drawn by Carl Semper during his voyage to the Philippines, and it was first named by Bergh (1875: pi. 41, fig. 10) based on Semper's figure. Later. Bergh (1890) published a complete descrip- tion based on additional specimens from Mauritius. This species has been also recorded in the Eastern Pacific Ocean from the Gulf of California to Panama (Marcus & Marcus, 1967; Houston, 1990; personal observation); in the Eniwetok Atoll (Marcus & Marcus, 1967); in north-


Notes, Information & News Page 405 Figure 1 Specimen of Titiscania limacina from Puerto Ayora, Santa Cruz, Galapagos Islands (17 mm in length). east Australia (Burn. 1975); and in the Molucas (Strack, 1998). Taki (1955) described the second species of this genus in Japan, Titiscania shinkishihataii, which was rediscovered and redescribed by Saito & Tsuchiya (1990). The differences between both species might not be considered specific characters because some minor differences have also been found in specimens from distant geographic areas, perhaps as a result of their different stage of conservation or as a consequence of the methods used to study the slugs. Therefore, both taxa might be synonyms. Nevertheless, according to Scott & Kenny (1998), the Titiscanidae remains a poorly known group with few specimens available for study. Titiscania limacina, despite being a shallow-water gastropod recorded from the west American continental coast and from some eastern Pacific islands, is not known from any of the eastern Pacific oceanic islands, as pointed out by Emerson (1991) in his study on the tropical trans-Pacific prosobranch gastropods. The malacofauna of the Galapagos Islands is, without doubt, the best studied of all eastern Pacific islands (see the reviews of Finet, 1991, 1994). However, Titiscania limacina has never been recorded there until now. During a Spanish scientific trip to the Galapagos Islands in 1991, organized by the Museo de Ciencias Naturales de Tenerife, Canary Islands, three specimens of this species were found. These specimens were collected off Puerto Ayora, Santa Cruz Island, in front of the Charles Darwin Research Station (6 March 1991). The three specimens were found beneath intertidal rocks, covered by white compound ascidians of the family Didemnidae. They measured alive 11, 14, and 17 mm when crawling. At resting position they were notably shorter. The external appearance of our specimens (Figure 1 ) does not differ from other descriptions of the species. These elongate sluglike animals were white in color with a row of 1012 opaque white papillae along each side of the notum. A white defensive secretion was expelled from these papillae when the animals were disturbed. Some isolated papillae were also present on the notum. The cephalic tentacles were long, thin, and sharply tapering. They became abruptly narrower just at their bases. Small eyes were located at the outer bases. The ctenidium projected partially from the opening of the mantle cavity near the head and somewhat toward the right side. The foot was translucent and wide, extending well beyond the notum, with its anterior end expanded into a pedal veil. The three specimens are deposited at the Museo de Ciencias Naturales de Tenerife, Canary Islands. Acknowledgments. We are indebted to Juan J. Bacallado, head of the project "Galapagos: Patrimonio de la Humanidad" for inviting us to participate in the scientific trips to the Galapagos Islands. Ignacio Diez Cortaberna made the definitive drawing based on the original sketches of the authors. Literature Cited Bergh, R. 1875. Malacologische Untersuchungen. Pp. 315-376, pis. 40-48 in C. Semper, Reisen im Archipel der Philippinen. Zweiter Theil. Wissenschaftliche Resultate nos. 8 & 9. Bergh, R. 1890. Die Titiscanien, eine Familie der rhipidoglossen Gastropods. Morphologisches Jahrbuch 16l)l-26, pis. 1-3. Burn, R. 1975. Titiscania limacina Bergh, 1875, an unusual gastropod new to Australia. Australian Shell News. 111. Emerson, W. K. 1991. First records for Cymatiwn muudiim (Gould) in the Eastern Pacific Ocean, with comments on the Zoogeography of the Tropical Trans-pacific Tonnancean and non-tonnacean prosobranch gastropods with Indo-pacific faunal affinities in West American Waters. The Nautilus 1052)62-80. Finet, Y. 1991. The marine mollusks of the Galapagos Islands. Pp. 253-280 in M. James (ed), Galapagos Marine Invertebrates. Plenum Press: New York. Finet, Y. 1994. The Marine Mollusks of the Galapagos Islands: A Documented Faunal List. Museum dhistoire Naturelle: Geneve. 180 pp. GOSLINER, T. M, D. W. Behrens & G. C. Williams, 1996. Coral Reef Animals of the Indo-pacific. Sea Challenger: Monterey. California, 314 pp. Houston, R. S. 1990. Reproductive systems of neritimorph ar-


surface of the gill skeleton in primitive Wrasses. The varied colors of the Wrasses and Parrot Fishes are due to the high development of their chromatophores or pigment-bearing organs. Sculpins, Gurnards, Scorpion Fishes, etc. (Suborder Scorpaenoidei): This group is generally known as the cheek-armored fishes because of the fact that one of the bones surrounding the eye is much enlarged and arches backward over the cheek so as to gain a broad contact with the forepart of the bony gill cover. Thus it forms the so-called "bony stay" that strengthens the skull in this group. The Gurnards, or Sea Robins, with their broad heads enclosed in this bony shield, bear little resemblance to the Basses, yet they are connected with the primitive Sea Bass stock through the Sculpins and Rosefishes. Most of the fishes of this group are marine forms, living either among the rocks or on or near the bottom, sometimes at considerable depths. The most primitive forms are the Rosefishes; the most advanced are the Gurnards. The first three rays of the Gurnards' breast fins are specialized as "legs" These rays are separated from the rest of the fin and from each other, and due to this specialization the movements of the fish among seaweed or along any surface much resembles walking on these fins. In the Lumpfishes and the Sea Snails, the ventral fins are united and form a sucking disk by means of which these clumsy, slow-moving fishes cling to rocks. The Platycephalids are a group of fishes from the East Indies and Japan. As their name implies, they have flattened heads, which present a curious resemblance to the heads of bottom-living forms of widely different groups, such as the extinct Ostracoderms (see Fossil Fishes, page 82) and the Armored Catfishes (page 38). The Gobies (Suborder Gobioidei): The Gobies have pushed their way into all the seas outside the Arctic and Antarctic circles, and have representatives in the fresh waters of all parts of the world. The central form, Gobius, is not very different from the Johnny Darters of the Perch-bass group, except that the ventral fins tend to form a sucking disc by means of which the fish can cling to rocks. In the Mudskipper, Periophthalmus, the pectoral fins are modified and strengthened so that they can be used as flippers, and the fish skips about on these over the mud flats of eastern tropical rivers. Its eyes are greatly enlarged and protruding. In the Blind Goby of California, on the other hand, the eyes are reduced to mere vestiges, and the fish lives like a slug under rocks. In size, the Gobies vary from the minute Pandaka pygmaea of the Philippines, which measures only 10 millimeters (less than inch) in length, to Eleotris marmorata of Siam, which grows to nearly 3 feet. The Gobies differ from the cheek-armored group in lacking the bony stay of the cheek. Blennies (Suborder Blennioidei) The Blennies are active little fishes living around rocky or coral reef shores. Certain Blennies, for instance the Lizard Skipper of Samoa, even leap from rock to rock at low water, in this respect paralleling the unrelated Mudskipper. Most of the Blennies are small, but there is one very large marine member of the family. This is the Sea Wolf, Anarrhichas lupus. This vicious fish has a long body and strong, tusklike front teeth, and is very hard to handle if caught. It also has large, rounded, crushing teeth on the roof 56


No. 508] NOTES AND LITERATURE 255 Knrraehee. Tn 1890 Dr. Hartlaub described Dichrometra Munzingeri from Koseir, and identified the Comatula leucomelas of Riippel with the Alccto pahnata of Professor Miiller. Mr. Chadwick did not find Tropin in, carinata nor Dichrodid find the other two species known from the Red Sea. Tlctcrometra savignii and Dichrometra panama; the remaining four species recorded include one family and three genera new to the Sfrphanom, Ira marginata and Dichrometra protcctus. Oligorncira scrripinna is recorded from Suez Bay. where it was dvdii-d M a deptli of Id fathom, the specimens differ from the type in the greater number of cirrus joints, and. in view of the general constancy of the cirrus characters in this genus, may eventually turn out to be a recognizable form, as may also those recorded under the same name from Ceylon; correlated, as usual, with the more numerous cirrus joints, the lower pinnules have also more numerous joints. Iridometra parvicirra, discovered by the "Challenger" among the Philippine Islands, is recorded from ten fathoms in Suez Bay. The single specimen has a large number of cirrus joints for the -onus, which, together with the furrowed first primibraeh. would suggest that it was rather closer to the . nana group of species than to I. parvicirra; the former occur from Mauritius to Japan, while the latter were previously known from the Philippines and Japan. We share the author's doubt in regard to his identification of Stephanometra marginata, which he records from Suez Bay, in ten fathoms. Dichrometra protectus (under the later name in, pari pinna) is given from Suez Bay, and Sikkim. The number of arms m the specimens is unusually small, and it would have been well worth while to have recorded their size. In addition to the localities cited by Mr. Chadwick, the species is known from Cebu, Philippines, Fiji and Singapore. Dichrometra palmata was found on the coral reef at Misharif Island, Khor Dongola, and from between tide marks at Suez. Heterometra savignii (emended, following Carpenter, to savignyi) is recorded from four fathoms in Suez Bay, from nine fathoms at Ul Shubuk, from ten to twelve fathoms at Khor Shinab, and at the anchorage at Salaka; the distribution of the species is given as "Red Sea, Ceylon" hut I am unable to recall any record from the latter place ; Carpenter gives it from


Plate 2402, SHOREA RIGIDA, Bmndis. DiPTEROCARPACEAE. Tribe Shoreae. S. Ti^i^^ Brmidis (sp. nov.) ; arbor ingens, ramuHs petioHsque verrucoso-scabris, novellis steJlato-tomentosis, foliis rigide coriacois iiiargiue revolutis elliptico-oblongis supra glaberrimis nitidis infra l)]Jis stellatis hirsutis nervis secundariis utrinque 18 arcuatis subtus valde prominentibiis nervis tertiariis parallels, floribus sessilibus in pamculis axillanbus et terminalibus, ebracteatis, sepalis injequaliljus extus pilis longmsculis hirsutis intus ad apicem puberulis, petalis oblongis obtusis apice barbatis, staminibus circiter 50, filamentis basi connatis petahsque adhairentibus antheras ter vel qnater superaiitibus anthens fere orbicularibus omnino exappendiculatis, ovario inferne glabro, superne in stylopodiuni elongato-conicum pilis longiusculis tomentosum angustato, stylo teuui glabro stylopodium vix cTquante stigmate lato obtuse 3-dentato, fructu tomentoso acuminato basi tantum calyeis segmentis tecto. Hab. Singapore (garden jungle), Ridley (No, 6393). f poll longa ad 2 poll lata; petiolus 3/4 poll longus. Calycis iructiteri segmenta tria majora lineari-oblonga, obtusa, basi angustata, lU-nervia, 4-5 poll, longa. Fructus % poll, longus. Cotuledonea crass:«, carnosae, major bifida ; radicula brevis. Attains 180 feet, with a tall clean stem. Foliage light green ; flowers creamy white in great masses ; wings of fruiting calyx reddish brown. Flowers in July. This species belongs to the small section of Shorea with inappendiculate anthers which now comprises 6 species of the Malay Peninsula, Sumatra, and Borneo. Some of these, like S., leprosula, Miq., have 15 stamens, while others (S. sericea, riyida) have a large number. D. Brandis, Tig. 1. Portion of leaf showing stellate indumentum. 2. Expanded flower. ^-Same enlarged. 4. Stamen. 6. Vertical section of ovary. Except fig 2. enlarged.


INTRODUCTION 4 THE FIRST EDITION OF BLANCOS FLORA DE FILIPINAS The first edition of Blanco's Flora de Filipinas was printed in Manila in 1837. It is an octavo volume consisting of a prologue, a general introduction, and the systematic arrangement and description of about 903 species under binomial names. The work is written wholly in Spanish. The prologue briefly relates the circumstances under which the work was prepared and contains a review of the botanical work accomplished by Blanco's predecessors in the Philippines; namely, Fathers Clain, Delgado, Mercado, and Santa Maria, but with no mention of the vastly more important work of Camel; and a brief mention of the work of Nee and Pineda, of the Malaspina Expedition, but no mention of the more important explorations of Haenke, of the same expedition. The general introduction, occupying pages IX-LXXVIII, consists of an explanation of the terms used in descriptive botany and an explanation of the Linnean system of classification, with a list of the admitted genera arranged under their Linnean classes. The generic descriptions are greatly: abbreviated, but the specific descriptions are usually long and detailed, although often general in nature; some of them are very short and imperfect-To these are often added long or short discussions, giving data of economic importance, occurrence, time of flowering, native names, etc. ; this supplementary data is often of more value in locating and interpreting Blanco's species than are the descriptions themselves. The work is supplied with an alphabetic index to generic names; one to native names, in which not all of the native names mentioned in the text are included; and a tabulation of the various genera according to the economic uses of the various species. In the text various species that Blanco was unable to refer to their proper generic positions are described under their native names. These are usually placed under the Linnean classes to which they pertain and cannot in any sense of the word be Flora de Filipinas. Segu Por el P. Fr. 1-7. 7


8 SPECIES BLANCOANAE treated as genera, nor were they so considered by Blanco. They include descriptions of: A lama. Apulong, Balihai, Banago. Binoicang. Bitlag, Calamansanay Calico Camotain, Corong. Cunalon, Octodon. Ge-jua. Hinguio, Iloilo. Inoyaban. Lanzones, Lapolapo. Loctong. Longayan. Malulucban, Oyisan. Palindan, Pandapanda. Pasac. Padang labuyo Putian. Sangumay, In the present work no special attempt has been made to locate these species, except in those cases where Blanco himself or other authors have later made them the bases of binomials. With few exceptions none of the descriptions have any bearing on questions of nomenclature. The new genera proposed, all but three of which have proved to be invalid, as indicated by the reductions here given, are: Azaola (Payna) Baling ayum Calogyne) Calms ( zstrebltts) Cobanga zcanscora), Enrila ( Ventilago) Lumanaja ( Homonoia) Lunasia, Malaisia. Mamboga ( Mitragyna) Manungala Sainadra) Palaquium. Quilamum ( z Crypteronia) Dichapetalum) 9 Qtiilesia ( Soala ( . Sulipa — Gardenia). Tala ( z Limnophila) Tayotum ( Geniostoma) FLORA The second edition of Blanco's Flora de Filipinas in plan same a v after Manila Blanco's death, which occurred April 1, 1845. In the publication of this work a subsidy of 500 pesos was granted economic de amigos pais In this edition about 1,132 species are described under Latin binomials, and 27 under their native names. Four species described in the first edition are excluded while 198 are added, making the total number of species dpsprihrl in THIS first and Flora de Filipinas, segun el Sistema sexual de Linneo. For el P. Fr. Manuel Blanco, Agustino Calzado. Segunda impression corregidor y aumentada per el mismo autor (1845) I-LIX -f 1-619. Blair, E. H, and Robertson, J. H. The Philippine Islands 1493-1898, 50 (1907) 69; 52 (1907) 312.


10 SPECIES BLANCOANAE tion through three bulky and unwieldy folio volumes without adding a single item to our knowledge of the Philippine flora. Unfortunately page references to the first and second editions are omitted. In consideration of the date at which it was printed, it is indeed a curious publication. The third edition follows the em at least half a century. from from himself eliminated from name for the one given in the second. The only praise that can be given in this glorified edition of Blanco's Flora de Filipinas is that the Latin translations made Blanco's descriptions more available to botanists generally; yet nearly forty years before, Walpers had published Latin translations of about 180 of Blanco's diagnoses of new species, which is the part of Blanco's work of most general interest to taxonomists. Two editions were issued, one the "edition de luzon" the other the "edition economic" The cheap edition differs from the edition de luxe in that it is printed on less expensive paper, and the plates are not colored. The plates differ further from those of the edition de luxe in that they are numbered. The cheap edition was sold at 125 dollars per fascicle in the Philippines, and 175 dollars outside of the Philippines; while the edition de luxe was sold at 225 dollars and 250 dollars respectively, the prices in Mexican silver. Each fascicle was advertised to consist of sixteen pages of text and six plates. The cheap edition was to consist of numbered volumes; but the edition de luxe was to be numbered, and the edition limited to 500 copies. The plan of numbering the volumes was apparently abandoned. means a rare one; but the cheap edition is exceedingly rare. I have seen but a single incomplete copy of the latter, consisting of a complete set of the text and about 140 plates. This copy is fourth volume Manila Filipinas is by far the most important part of the work. It consists of three separately paged parts, of which the third, the Novissima Appendix. is the. mili nnp nf rpnl nino 'Linnaea 16 (1842) Litt-bericht 1-68.


12 SPECIES BLANCOANAE authors. A large number of them refer to species that were unknown to Blanco or at least to those that were not described by him. A very high percentage of the species figured are common and well-known ones of wide geographic distribution and include many of the common weeds and the various cultivated species, including the commonly cultivated ornamentals, some of which were introduced into the Philippines after Blanco's time. No attempt was made to select the endemic species in the preparation of the plates nor to confine the illustrations to those species not previously figured by other authors; but about 80 of the total of 473 species figured or less than 17 per cent, represent endemic species. In general the illustrations are good, but a few are unrecognizable with certainty beyond the genus. In some cases the color selections are very faulty. The detail draw-gs are usually msutticient, and many are p In order to make the present work more com to the Flora de Filipinas, I give below a list of the plates of the third edition that do not pertain to species actually deforms listed their species. Fernandez Bentham and Hooker arrangement stem LIST OF PLATES IN THE THIRD EDITION OF BLANCOS FLORA DE FILIPINAS THAT REPRESENT SPECIES NOT DESCRIBED BY BLANCO OR BY LLANOS 344 Dillenia reifferscheidia F-VIU. 148 Talauma villarii Rolfe (T. mutabilis Naves, non Blume), 198 Talauma coco (Lour) Merr, T. pumila Blume). 193 Phaeanthus suberosus Hook. f. & Th, (P. malabaricus Naves, non Bedd). 209 Capparis sepiaria Linn. 94 Dianthus chinensis Linn. 241 Calophyllum soulattri Burm, f, (Calophyllum spectabile WIWD) (poor). 333 Bombycidendron vidalianum Merr. & Rolfe Hibiscus vidalianus Naves) J4C Hibiscus syriacus Linn. (poor). u assigned utilized when they are the correct ones. In other cases, where Naves's wrong original name in parenthesis. with


INTRODUCTION 15 perdix nd it is wholly the work of Father Celestino Fernandez-villar and Father Andres Naves, but mostly the actual work of the former. Fernandez-villar is the author of the treatment of the dicotyledonous and gymnospermous orders, pages 1 to 212, and of the monocotyledonous orders and vascular cryptogams, from Fimbristylis bispicata on page 307 to the end of the work. Naves is the author of the monocotyledonous orders from page 213 to Fimbristylis nutans on page 307. The date appearing on the title page is 1880, and the introduction is dated December 12, 1880. The printer's date for the last part, indicated on page 375, is June 15, 1883. From an examination of an unbound copy in which the original fascicle covers were preserved I find that pages 1 to 272 were issued in 1880; pages 273 to 336 were issued in 1882; and pages 337 to 375 were issued in 1883 As indicated in the introduction to the third edition of the Flora de Filipinas it was the intention of the authors that de bonn Mercado Blanco, and Llanos; all of those described from Philippine im material V 77arranged in their natural orders. prospectus, issued in 1877, is that descriptions would be added, but this is not definitely stated. However, owing to various circumstances the authors were obliged to abandon their original plan in part, as indicated in the introduction to the Novissima Appendix The reasons given were the enervating effect of the climate, the impatience of the majority of the subscribers for the termination of the work, the lack of special training part of tl botanical botanical material preserved means European and American herbaria and had not seen a single specimen of the large Cuming collection, on which up to that date most of the actual knowledge of the Philinninp flint-a rn 12 Naves, A. and Fernandez-villar, C. Novissima Appendix ad Floram Philippinarum R. P. Fr. Emmanuelis Blanco, seu enumeratio contracta plantarum philippinensium hucusque cognitarum. Cum synonymis P.P. Blanco, Llanos, Mercado et aliorum auctorum. (1880-1883) IX -f 1-375. Merrill, E. D. The dates of publication of the third edition of Blanco's Flora de Filipinas. Philip, Joum. Set. 12 (1917) Bot. 113-116. " 1 (1877) IX, X. "Novis. App. (1880) VIX.


254 p Jp WHITEHEAD RANGE Suez Canal, Red Sea (Gulf of Suez), Gulf of Aden (Mukalla, Qishm, Aden), Gulf of Oman, but not recorded from the Persian Gulf ; elsewhere, Indian Ocean from Mauritius to East Indies, and from China, Australia, Micronesia, Polynesia. Specimens 6 fishes, 97-114 mm, Mukalla (B.M.N.H. 1962326102-107). 6 fishes, 99-115 mm, Mukalla (B.M.N.H. 1962326111-116). 3 fishes, 54-58 mm, Aden (B.M.N.H. 1962326108-110). 1 fish, 119 mm, Gulf of Oman (Z.M.C. Ci Blegvad's S. sindensis). 2 fishes, 118-119 mm, Gischin, coll. Hein (N.M.V. 1260). Sardinella fimbriata (Valenciennes) Sefer, Hashinch, Moomagh (Iran) Spratella fimbriata Valenciennes, 1847, Hist. Nat. Poiss, 20 (263) 359, pi. 601 (Type locality Malabar) Sardinella fimbriata Blegvad, 1944, Danish Sci. Invest. Iran, pt. 3 65, fig. 29 (copied Day) (off Fort Dilam) ; Fowler, 1956, Fishes of the Red Sea, 1 66 (on Philippine specimens). Note on synonymy. Bertin (19440), after an examination of the Valenciennes types of Clupeonia jussieui and Spratella fimbriata, concluded that the two represented a single species, which he believed to have " environ 70 branchiospines a la branche inferieure du premier arc branchial ". But, until the validity and identity of Lacepede's C. jussieu can be definitely confirmed or rejected, it seems best to retain the name fimbriata for the material described here. S. jussieu (as understood here, ie. sensu Fowler, 1941, 1956) very closely resembles S. fimbriata. Dutt (i96ia, 1962), who studied the two species off the Waltair Coast of India, stated that S. fimbriata makes its appearance there earlier than 5. gibbosa (ie. S. jussieu), but that at times when the two species coincide, their shoals are almost always discrete (Dutt 1962). He distinguished the two species on gillraker counts which, although increasing slightly with size of fish, remain just distinct (a difference of 2, 2, 3, 6, 6, 5, 8, 5, 2, 4, o, 4 rakers respectively in the 10 mm. groups 30-140 mm. standard length). In my material (24 fishes, 87-118 mm) I have assigned specimens with a count of 53-58 rakers to S. jussieu, and those with 60-72 rakers to S. fimbriata. In this size range, Butt's ranges in gillraker counts are respectively 43-60 and 57-77, with average values of 48, 52, 53, 56 for S. jussieu and 63, 66, 69, 70 for S. fimbriata in the 10 mm. groups represented (mean values of 54-9 and 66-5 respectively in my material) In the specimens examined here, there are no other meristic differences between the two species, and proportional measurements are almost identical except for a slightly shorter snout in S. jussieu (6-4-7-3 per cent, of S.L. ; cf. (6-7) 7-3-8-0 in S. fimbriata). Fowler (1956) finds a difference in numbers of scales in lateral series, S. fimbriata having a lower count (36-38 ; cf. 40-42), but counts are not always possible or perhaps accurate in these fishes.


DELFINADO: PHILIPPINE BITING MIDGES 651 base, tibiae with narrow basal band; hind tibial comb with four spines, the one nearest the spur longest. Wing (pi. 9, fig. 1) with costa extending 063 (054-070, w-10) of wing length; macrotrichia long and dense on distal half of cell R 5 sparse on cells Mi and Ms. Wing pattern as figured; extreme tip of second radial cell included in a pale spot; narrow pale spot at wing base; two prominent, broad, transverse pale bands across wing, the first at level of crossvein r-m to tip of anal cell, and narrowly interrupted by dark stem of mediocubital fork, the second beginning on and including tip of second radial cell and extending to tip of cell M 4 ; lighter pale spot at margin of wing on cell M 2 absent on cells R 6 and Mi. Halter brownish. Spermathecae (pi. 13, fig. 10) two, equal, pyriform, each measuring 0040 by 0054 mm; only rudimentary spermatheca present, ring absent. Male genitalia (pi. 14, fig. 4). Ninth sternum broad, with deep mesal excavation, the ventral membrane hairy; ninth tergum short and tapering, with long apicolateral processes, the apical margin between them with a deep, median notch. Basistyle short, mesal margin simple, only the dorsal roots developed; dististyle nearly straight, slender. Aedeagus with heavily sclerotized basal arch, width at base 07 times total length, anterior membrane covering about distal half of arch; distal stem short, the apex broadly rounded, unsclerotized. Parameres with main bodies joined at bases by a thin loop; knobbed base straight, with stout, short, divergent stem and pointed, bare tip. Holotype. A female, Clark Air Base, Angeles, Pampanga, September 17, 1957, light trap, I. Balatbat. In the collection of the Division of Malaria, Philippine Department of Health. Allotype. A male, same data as holotype (PDH). Paratypes. Seven females, same data as holotype (3, PDH; 3, USNM; 1, BPBM); 1 female, Tala, Rizal, May 22, 1958, light trap, M. D. Delfinado (BMNH). Plesiotypes. One female, same data as holotype (PDH); 1 female, CNHM-11, Dimanianga, Busuanga Island, Palawan, near sea level, H. Hoogstraal (CNHM). Culicoides (Oecacta) perornatus, new species The characteristic distal pale spot on cell R 5 the remarkably swollen third palpal segment with large, deep sensory pit, and the elongate-oval spermathecae with long, chitinized neck are the distinctive features of this species. Female. Length about 16 mm, wing 10 mm. long. Eyes contiguous for the length of the diameter of 35 facets (pi. 12, fig. 9). Antennae missing. Palpal segments in proportion of 2037612337; third segment greatly swollen (pi. 11, fig. 11), 20 times as long as broad with a large, deep sensory pit at middle. Mandible teeth 10. Legs brown; knees with a narrow darker brown band, with adjacent yellowish band on front and middle tibiae; fore and mid femora banded; base and apex of hind tibia pale, tibial comb with four spines. Wing (pi. 9, fig. 2) with costa extending 063 of wing length; macrotrichia fairly abundant over distal half of


DELFINADO: PHILIPPINE BITING MIDGES 663 Halter brownish. Spermathecae (pi. 13, fig. 17) two, ovoid, unequal, measuring 0034 by 0051 and 0034 by 0047 mm; rudimentary spermatheca and ring both present. Male genitalia (pi. 14, fig. 5). Ninth sternum broad, with a deep median excavation, the ventral membrane bare; ninth tergum rounded apically with a well-developed, thinly sclerotized median lobe, the apicolateral processes absent. Basistyle stout with setae on mesal margin, poorly developed ventral roots, short, pointed dorsal roots; dististyle slender, slightly curved to a blunt apex. Aedeagus long and narrow, width at base 04 times the total length, the basal arms well sclerotized and stout, a sclerotized anterior band (transverse bar) near base of arch, distal peg absent but with thinly sclerotized membrane forming a small distal arch; apex long and slender with a ball-like tip. Parameres separate, base abruptly bent and slender; stem nearly straight, tapering to recurved haired tip. Holotype. A female, CNHM-6, Burungkot, Upi, Cotabato, 1500 feet alt, January 1-6, 1947, net along forest floor, F. G. Werner. In the collection of Chicago Natural History Museum. Allotype. A male, same data and depository as holotype. Paratypes. A male and female, same data as holotype (USNM). Culicoides (Culicoides) malaya Macfie Culicoides malaya Macfie, 1937, Ann. Trop. Med. Parasit, 31 (4), p. 471 (female; Malaya; wing illus). Diagnostic characters. Wing with second radial cell in a pale spot, pale markings extensive, distal pale spots on cells R 5 Mi, and M 2 not touching margin of wing, cell M 4 dark at base, the distal pale spot broadly meeting margin of wing (pi. 10, fig. 6); eyes contiguous; third palpal segment moderately swollen, with small, shallow sensory pit; hind femur entirely brown, tibiae with narrow, yellowish band at base and apex; two spermathecae well sclerotized, suboval; male aedeagus with anterior marginal band, distal peg, truncated tip; parameres separate, tips bare; basistyle with ventral roots not developed; ninth tergum with truncated apex, very small apicolateral processes. Distribution. Mindanao: Pikit, Cotabato, December 16, 1946, at light, F. G. Werner. Culicoides (Culicoides) recurvus, new species The male genitalia of recurvus are nearly identical with those of hirtipennis to which it is closely related, but recurvus may be distinguished by vein R 4+5 which is dark up to the point where it turns abruptly forward to meet the costa; also, the front knee is dark, with an adjacent pale band on tibia. Female. Length about 15 mm, wing 103 (099-106, n-8) mm. long. Eyes widely separated (pi. 12, fig. 17), bare. Antenna with flagellar segments in proportion of 443034343434343751546164102, antennal ratio 117 (114-120,


664 FIELDIANA: ZOOLOGY, VOLUME 33 n-3); antennal sensoria present on segments III, VII, XI-XV and III, XI-XV. Palpal segments in proportion of 1744612730; third segment (pi. 11, fig. 19) greatly swollen, 20 times as long as broad, with a large, shallow, sensory pit. Mandible teeth 15 (13-19, n-8). Legs brown; front knee dark with adjacent pale ring on tibia; femora dark; bases of middle and hind tibiae yellowish; hind tibial comb with five spines, the second from the spur longest. Wing (pi. 10, fig. 7) with costa extending 064 (061-067, -8) of wing length; nearly bare with only a few macrotrichia at wing tip. Wing brown with limited rounded pale spots appearing yellowish; distal end of second radial cell pale; pale spot over crossvein r-m separated anteriorly from costal pale spot by the radial vein; cell R 5 with distal pale spot before margin of wing; cells M 2 and M 4 each with submarginal pale spots, cell M 2 also with a double pale spot from just below medial fork to anterior level of mediocubital fork; tip of vein Mi distinctly pale. Halter brown. Spermathecae (pi. 13, fig. 18) two, subspherical, subequal, measuring 0037 by 0051 and 0040 by 0051 mm; rudimentary spermatheca and ring both present. Male genitalia (pi. 14, fig. 7). Ninth sternum broad, with a shallow median excavation, the ventral membrane bare; ninth tergum rounded apically with a thinly sclerotized median lobe, the apicolateral processes absent. Basistyle slender, mesal margin hairy, only the dorsal roots developed; dististyle thick at base, narrowing toward apex. Aedeagus long and narrow with anterior sclerotized band (transverse bar) near extreme base of arch, very long and slender tip, distal peg absent. Parameres separate, with abruptly bent, small, knobbed base, slender and straight stem, and recurved, pointed, and bare tip. Holotype. A female, Clark Air Base, Angeles, Pampanga, September 17, 1957, light trap, I. Balatbat. In the collection of the Division of Malaria, Philippine Department of Health, Manila. Allotype. A male, same data and depository as the holotype. Paratypes. Two females, same data as the holotype (1, CNHM; 1, USNM) ; 1 female, Taft, Samar, May 3, 1956, carabao-baited trap, I. Balatbat (USNM); 1 female, Davao, May, 1956, carabao-baited trap, H. Crisostomo (PDH). Culicoides (Culicoides) sumatra Macfie Culicoides sumatra Macfie, 1934, Tijdschr. Ent, 77: 215 (male; Sumatra; genitalia illus); Ann. Trop. Med. Parasit, 28 (2), p. 190 (1934) (female; Malaya, at light). Diagnostic characters. Wing with distal half of second radial cell in a pale spot, transverse pale spot at middle of cell R 6 reaching anterior margin of wing, pale tip of vein Mi, dark base of cell M 4 (pi. 10, fig. 8); eyes contiguous; antenna with distal sensory tufts present on segments III, XI-XV; third palpal segment swollen, with small round sensory pit; two well-developed spermathecae; male aedeagus with anterior marginal band, distal peg, and spherical tip; parameres separate, tips bare; basistyle with ventral roots not developed; ninth tergum rounded apically, apicolateral processes absent.


F I MAY 3 1 1! ZOOLOGICAL SERIESJJNIVERSITY OF ILL1NOI! OF FIELD MUSEUM OF NATURAL HISTORY Volume XX CHICAGO, MAY 15, 1935 No. 8 A NEW CROCODILE FROM THE PHILIPPINE ISLANDS BY KARL P. SCHMIDT ASSISTANT CURATOR OF AMPHIBIANS AND REPTILES The salt-water crocodile, Crocodylus porosus, which is widespread in the East Indies, is the common crocodile of the Philippine Islands, ranging throughout the archipelago. Very large specimens occur, and the largest skull on record, in the Museum of Comparative Zoology, was traced to its source in Luzon by Barbour (1924, p. 16). Two large skulls in Field Museum, collected in Lake Naujan, Mindoro Island, in 1891, by D. C. Worcester and F. S. Bourns for the Menage Expedition, unquestionably represent the species porosus. These attracted my interest and were cleaned and examined critically in drawing up the description of Crocodylus novae-guineae in 1928. During a short stay in the Philippines en route home from the Crane Pacific Expedition of Field Museum, in 1929, I obtained a small crocodile of the species porosus at Zamboanga, Mindanao. During this expedition, a series of skulls of both porosus and novae-guineae was obtained in New Guinea, and my interest in these species renewed. On arriving in Manila, accompanied by Mr. Walter A. Weber, artist of the expedition, we were most cordially received by Mr. W. H. Brown, Director of the Bureau of Science, and Mr. H. R. Montalban, of the Division of Fisheries. Through the interest of these gentlemen, three small crocodiles, presumably also from the vicinity of Lake Naujan, were obtained for us from the island of Mindoro and presented to Field Museum. These specimens were unfortunately long delayed in transit, so that they reached Chicago as dried mummies. Greatly to my surprise, these Mindoro specimens proved to be entirely distinct from the species already at hand from Lake Naujan, and to be equally different from Crocodylus palustris, recorded from Borneo and Palawan, as well as from C. siamensis of southeastern Asia. Further examination of this material was postponed in the hope that additional specimens might be obtained. In examining some uncleaned crocodile heads (without data) stored in Field No. 337 67


68 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XX Museum, I find one which exhibits the six postoccipital scutes of the Mindoro form, and on preparation of this skull it proves to agree so minutely with the three smaller skulls cleaned up from the dried specimens that there can be no doubt that it is also from the Philippines, received either with the Menage Expedition collections, or from Dr. J. B. Steere, who is also known to have collected on Mindoro. The examination of the four specimens available convinces me that the species is certainly distinct and undescribed, and I feel that its publication will call attention to its existence and promote further collecting. It may be named Crocodylus mindorensis sp. nov. Type from the island of Mindoro, Philippine Islands. No. 11135 Field Museum of Natural History. Received in 1929 from the Philippine Bureau of Science. Mummified body and cleaned skull. Diagnosis. Allied more closely to Crocodylus novae-guineae than to any other species, agreeing with it in the characters which distinguish it from C. porosus; namely, presence of occipital scutes, more incised webs on hind feet, enlarged pectoral plates, scutes of dorsal shield in contact laterally, inferior opening of internal nares, and uninflated palatines. Distinguished from novae-guineae by its proportionately broader and heavier skull, larger eustachian tube, palatines less parallel-sided, heavier pitting of skull and more pronounced antorbital ridges. Description of type. Dorsal shield consisting of 17 transverse rows of scutes, composed of 8 scutes each at mid-body, the bony scutes in contact; postoccipitals 6; nuchals 4-2-2; web" of hind toes strongly incised; a row of enlarged pectoral scutes, behind which there are 25 transverse rows of ventral plates to the anterior border of the anus; 18 segments of the tail with a double crest, followed by 18 single-crested segments. Skull of intermediate type, neither extremely broad nor elongated. Snout distinctly broader than in either porosus or novae-guineae. Antorbital ridge high and abrupt laterally as compared with novae-guineae, failing to continue on the naso-maxillary suture as it does in porosus. Pitting of the dorsal surface much deeper and coarser than in skulls of the same size of either novae-guineae or porosus. Premaxillary teeth 5; festooning of maxillary teeth pronounced, more so than in the allied species; opening of internal nares downwardly directed; eustachian opening of skull larger than in novaeguineae, about as in porosus; form of cranial table intermediate between that of porosus and novae-guineae; posterior border of frontal


is 70 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XX (less than 6 feet). The correspondence of this skull, No. 19891, even in minor details, with the three subsequently received from Mindoro is extraordinary. Remarks. The new Philippine species is quite obviously to be interpreted as a fresh-water species, and where porosus enters inland lakes it is doubtless found in marshes and smaller bodies of water. Malcolm Smith (1919, p. 218; 1931, p. 42), in distinguishing C. siamensis from C. porosus calls attention to the juxtaposed elements of the dorsal shield, the enlarged pectoral plates, and the more incised webs of siamensis. Crocodylus mindorensis and novae-guineae agree with siamensis in these respects. The skull of mindorensis is much more slender and lighter in construction than that of palustris. It differs from siamensis in the narrow interorbital space, the five premaxillary teeth, the absence of a median ridge on the frontal, and the more posterior position of the enlarged fifth maxillary teeth. The new form does, however, form a link in the chain of species of fresh-water crocodiles with mutually exclusive ranges palustris in India, siamensis in southeastern Asia, mindorensis in Mindoro, novae-guineae in New Guinea, and johnstoni in Queensland. I suspect that C. biporcatus raninus of Miiller and Schlegel may be the Bornean representative of this fresh-water series, and it is reasonable to suspect another undescribed species in the ancient lakes of Celebes. Field observation of the habitat relations of Crocodylus mindorensis with C. porosus affords an interesting problem for any naturalist visiting the Philippines. I have followed Stejneger (1933, p. 117) in retaining Laurenti's generic name Crocodylus in the long customary sense. REFERENCES BARBOUR, THOMAS 1924. An Historic Crocodile Skull. Copeia, 1924, p. 16. SCHMIDT, KARL P. 1932. Notes on New Guinean Crocodiles. Field Mus. Nat. Hist, Zool. Ser, 18, pp. 165-172. SMITH, MALCOLM A. 1919. Crocodilus siamensis. Journ. Nat. Hist. Soc. Siam, 3, pp. 217-221, pis. 4-5. 1931. Reptilia and Amphibia. Vol. I Loricata, Testudines (Fauna of British India). Taylor and Francis, London, 8vo, xxvi 185 pp, 41 text figs, 2 pis. STEJNEGER, LEONHARD 1933. Crocodilian Nomenclature. Copeia, 1933, pp. 117-120. WORCESTER, DEAN C. 1898. The Philippine Islands and Their People. Macmillan Co, New York, 8vo, XX 529 pp, illus.


162 FIELDIANA: ZOOLOGY, VOLUME 51 H. a. catarmanensis Camiguin South cf 70-75 9 71-76 mm. Range. — Camiguin South; lowlands to 4,950 feet on peak of Mount Camiguin. Remarks. — For comparison we have large series of H. a. azurae from Luzon, Negros, and Mindanao, moderate series from Palawan, Bohol, Siquijor, Samar, Cebu, and one or a few specimens from each of Polillo, Sibuyan, Romblon, Mindoro, Guimaras, and Basilan. Males and females are about equally represented in all these series, every blue-backed bird has the black nape patch, usually of considerably greater extent than in that of the Camiguin South birds with the largest patch. The size of the black throat band is variable, rarely absent. All the females have brown backs (of various shades of brown or greyish-brown) There is always a possibility of some mis-sexing of specimens in the field, but the blue-backed females from Camiguin South are not like any specimens, male or female, from any other islands. The following tentative conclusions are advanced. There is a reduction in sexual dimorphism through the female becoming partly cock-feathered, notably in the blue back, and the male becoming more like this new type female through the partial loss of the black nape patch. There is an indication, from this collection, that the female may be dimorphic, most (4 out of 5) are much like dull males but without black markings; a few (1 out of 5) are more like the female of H. a. azurae. However, this may be an earlier plumage, special to the immature female in this population. There are two other species of the genus in Philippines: H. helenae, Camiguin North, Luzon (series Field Museum), Samar, Mindanao; and H. coelestis, Luzon, Negros, Sibuyan, Dinagat, Mindanao, Basilan; and neither is known to occur on Camiguin South. One may be tempted to consider the possibility that this absence has allowed sexual dimorphism to decrease. But, there are other islands in the Philippines, with only H. azure; and over most of the range of H. azure it is the only representative of the genus. Birds from Nepal have the same male and female mode of plumage as do those of H. a. azurae of Philippines. Dicaeum trigonostigma Salomonsen, 1960, Amer. Mus. Novit, No. 1991, pp. 28-37.


RAND AND RABOR: BIRDS OF CAMIGUIN SOUTH 163 This is a widespread species presumably originating in the Philippines (though absent from Palawan) where Salomonsen recognized nine subspecies, most of them with striking and clear-cut characters. In all the rest of the range (Java and Borneo to Assam), only five subspecies have developed and most of them with rather trivial characters. Five of the nine subspecies recognized in the Philippines are small island subspecies: on Romblon and Tablas (intermedium); Sibuyan (sibuyanicum) ; Siquijor (besti); Jolo, Tawi-tawi, Siasi (assimile) ; and Sibutu (sibutuense) An additional subspecies proves to have been developed on Camiguin South, raising the number of Philippine subspecies to ten, and six of them small island subspecies. The near relatives of the Camiguin South form seem to be the geographically adjacent cinereigularae of Mindanao-samar-leyte-Bohol area, and besti of Siquijor, and it is with these that comparison is needed, differing as they do chiefly in color of chin and throat, and in size. Dicaeum trigonostigma besti Steere Diagnosis. — Male, chin and upper throat yellow; rest of throat and sides of neck uniform grey, without a yellow wash. Size, small. Male, wing (10), 50-53 (av. 518 mm); culmen (10), 13-14 (av. 133 mm). Dicaeum trigonostigma cinereigulare Tweeddale Diagnosis. — Male, chin and upper throat yellow; lower throat and side of neck grey washed with yellow. Size, small. wing culmen Samar, male (10) 50-52 (av. 51) (10) 11-135 (av. 124) Bohol, male (4) 49-52 (av. 50) (4) 12-13 (av. 125) Mindanao, male (10) 49-52 (av. 512) (10) 12-135 (av. 127) Dicaeum trigonostigma isidroi, new subspecies' Type— Field Museum of Natural History no. 284521 from Kasangsangan, Catarman, Camiguin South Island, Philippine Islands. Adult male collected June 11, 1968, by D. S. Rabor. Diagnosis— Like both of the above subspecies but differs in the male having the chin and upper throat much paler yellow; in having This new race is named in honor of Dr. Antonio Isidro, president of Mindanao State University, and in appreciation of his great interest in biological research.


164 FIELDIANA: ZOOLOGY, VOLUME 51 the lower throat pale, whitish grey (without any yellowish wash) becoming darker toward the sides of the throat; size larger. Wing cf (10), 55-57 (av. 556 mm); culmen (10), 135-15 (av. 14 mm). Range. — Camiguin South Island, 1,000-2,500 feet altitude. Remarks. — Most races of this species in Philippines have wing 48-53 mm; but there are two larger subspecies, the present one (see above) and sihuyanicum wing 54-57 mm, both small island species. Zosterops nigrorum Mees, 1957, Zool. Verh. Rijk. Mus. Nat. Hist. Leiden, No. 35, pp. 160-170; Mayr, 1967, Peter's Check-list of Birds of the World, 12, pp. 299, 300. This is the only species of Zosterops in the Philippines that has completely yellow underparts. It is a species of large and small islands: large islands in the north, and the western central islands — Luzon, Mindoro, Masbate, Panay, and Negros; a very few scattered small islands — Camiguin North (north of Luzon); and a few small islands in the central Philippines with southern outposts on Cagayancillo (west of Negros, in Sulu Sea) and Camiguin South in Mindanao Sea. The relationships of this species are not clear, but the general appearance of the various subspecies, and their differences from other species, in appearance and range, seem to indicate a natural group, forming an endemic Philippine species, as treated in recent revisions. Up until now the known ranges of the related everetti, with underparts partly grey or white as well as yellow, and nigrorum have been allopatric. However, on Camiguin South Rabor found both species, everetti at low altitudes and nigrorum at higher altitudes. This is the only locality where both species occur. (It may be recalled that the old records of Z. everetti siquijorensis for Negros are in error, surely due to a mis-identification of specimens of Z. montana pectoralis) Seven subspecies of Z. nigrorum are recognized in the following, and are arranged in three groups. Group I. — Four races, generally bright to moderately bright in coloration; eye ring complete; little or no black or dusky in side of head; size smaller; wing cf 50-56 mm. Zosterops nigrorum meyleri Mcgregor agfecamiguin North. Diagnosis. — A small race, separable from all other races by wide, complete eye ring, especially wide below the eye, according to Mees


RAND AND RABOR: BIRDS OF CAMIGUIN SOUTH 165 whose scant material was variable in color. His pen and ink drawing of head indicates a short dusky streak below white eye ring only. Mcgregor, 1908, Manual Philippine Birds, pp. 618-619, says this race is similar to richmondi but lacks the black under eye and has eye ring wider. Wing (4), 535-56 (av. 544); culmen (4), 128-138 (av. 132 mm). (Mees) no specimens seen. Zosterops nigrorum aureiloris Ogilvie Grant Range. — Northwestern Luzon, south certainly to Mountain and Abra Provinces and Benguet; farther south intergradation with the next two races to be expected; also on Mindoro; sea level to 5,000 feet. Diagnosis. — A small, brightly colored race, complete eye ring, with, at most, a dusky smudge below it ; local area and a narrow area on forehead golden yellow and sharply contrasting with rest of upper-parts; upperparts bright yellowish olive green; underparts bright yellow with flanks lightly tinged olive. Male, wing (10), 49-56 (av. 530); culmen (10), 11-135 (av. 123 mm). Remarks. — Series from Ilocos Norte, Abra, and Mountain provinces. Collected by Rabor, in Field Museum of Natural History. Zosterops nigrorum sierramadrensis new subspecies Type. — Field Museum of Natural History no. 259880 from Luzon Island, Cagayan Province, Gonzaga, Mt. Cagua (in the Sierra Madre), 2,000-4,000 feet altitude. Adult male collected April 30, 1960. Diagnosis. — Like aureiloris of northwestern Luzon, but upper-parts darker, less yellowish, forehead darker yellow usually with an ochraceous tinge; underparts deeper yellow on the throat, and with more of an olive wash on breast and flanks; bill slightly longer. Thus, a more intensely colored, darker race. Wing c (10), 50-54 (av. 512 mm); culmen (10), 12-14 (av. 131 mm). Range. — Northeastern Luzon in Cagayan Province. Presumably ranges southward to intergrade with the next race in central Luzon. Remarks. — Series from Cagayan Province in Field Museum of Natural History. This is an additional example of northwestern and northeastern Luzon birds representing different subspecies, as


SOLEM: PHILIPPINE SNAILS OF ENDODONTIDAE Fig. 2. Discocharopa werneri, n. sp, CNHM 57194. Paratype. a, top view; b, basal view; c, side view; d, diagram of microsculpture (greatly enlarged). Scale line mm. Specimen lost in handling. umbilicus; the species with apertural teeth all have spirally striate apices. Remarks. The generic position of D. werneri is uncertain. The combination of radially ribbed apex, toothed aperture, and rather narrow umbilicus suggested the Polynesian-new Zealand Thauma- todon-ptychodon complex, rather than the toothless, widely um-


The Distribution and Ecology of Mammals on Leyte, Bi I i ran. and Maripipi Islands, Philippines Eric A. Rickart Lawrence R. Heaney Paul D. Heideman Ruth C. B. Utzurrum Abstract The land mammal faunas of the Philippine islands of Leyte, Biliran, and Maripipi are described. Collectively, a total of 48 indigenous species are known to occur on the three islands: 45, 30, and 25 from Leyte, Biliran, and Maripipi, respectively. These include 1 insectivore, 1 dermopteran, 33 bats, 2 primates, 7 rodents, 2 carnivores, and 2 ungulates. All but three of these species occur on the large neighboring island of Mindanao. The exceptions include a widespread but rarely captured vespertilionid bat and two rodents that have sister-species on Mindanao. Among the remaining 45 species, 9 are restricted to the Mindanao faunal region, 10 are more widely distributed in the oceanic Philippines, and 26 have distributions that extend outside of the Philippines. One species from Biliran and two from Maripipi (all poorly known insectivorous bats) have not been recorded from Leyte. Otherwise the faunas of the two smaller islands are subsets of the Leyte fauna, which is, in turn, a subset of the Mindanao fauna. Data indicate that the present-day mammal faunas of these islands have been shaped primarily by events that occurred during the last 1 2,000 years. Results generally support predictions concerning elevational patterns in species richness and abundance (decreasing with elevation for small fruit bats, increasing with elevation for small non-volant species) and habitat associations of endemic versus non-endemic species (the former restricted to pristine or lightly disturbed habitats, the latter predominant in disturbed habitats). However, some of these patterns appear to break down on small land-bridge islands with relatively depauperate faunas. Most notably, commensal or widespread Asian species are abundant in undisturbed or lightly disturbed habitats when there are few endemic species present. Many of the results have direct implications for wildlife conservation in the Philippines. Patterns of deforestation render lowland species most vulnerable. Habitat associations of species indicate that disturbances to natural habitat have a disproportionately greater negative impact on the unique endemic portions of a local fauna. The depauperate nature of faunas on small land-bridge islands demonstrates that relatively large faunal reserves are required to provide long-term protection for many species. Furthermore, elevational range restrictions or special ecological requirements of some species necessitate reserves that encompass the full range of regional habitat diversity. Introduction Mindanao, principally because early work revealed many endemic species in these areas. As a The Philippine Islands support a diverse mam-result of this limited focus, the mammals of most mal fauna of about 1 70 species, many of which of the smaller Philippine islands remain poorly are endemic to the archipelago (Heaney, 1986, known. Recent studies on some small and medi1991a; Heaney et al, 1987). Most studies of Philum-sized islands in the archipelago have yielded ippine mammals have focused on the faunas of a wealth of information on patterns of mammalian the highlands of the two largest islands, Luzon and distribution (Heaney, 1986; Heaney et al, 1989, FIELDIANA: ZOOLOGY, N.S, NO. 72, AUGUST 31, 1993, PP. 1-62 1


1991; Heaney & Rickart, 1990) and have underscored the importance of such areas both as natural laboratories and as potential faunal reserves. Unfortunately, this unique fauna is now severely threatened by habitat destruction. Primary forest cover in the Philippines as a whole has been reduced from more than 80% of the total land area near the turn of the century to ca. 8% in 1988 (Myers, 1988), and the destruction continues at an alarming rate. The impact of deforestation on wildlife may be particularly severe on small islands, where population and community structures may be less robust. Forest destruction also poses a very direct threat to human welfare. In addition to the loss of invaluable forest resources, the elimination of natural watershed protection in upland regions greatly increases the likelihood of seasonal flooding. On 5 November 1991, this last possibility became a terrible reality in northern Leyte, where thousands of Filipinos lost their lives in catastrophic floods and landslides associated with a tropical storm. The scope of this disaster was certainly magnified by the extensive logging that has virtually eliminated interior forests throughout most of the region. In this paper we present the first synoptic lists of mammals from three islands in the central Visayan group: Leyte, the eighth largest island in the Philippine archipelago, and Biliran and Maripipi, two small islands immediately north of Leyte (fig. 1). All three were part of the island of Greater Mindanao during a period of lower sea level in the late Pleistocene, becoming isolated about 10,000 years ago (Heaney, 1986). In relation to present-day Mindanao (with an area of 99,078 km 2 , Leyte (7,213 km 2 , Biliran (498 km 2 , and Maripipi (22 km 2 ) represent a graded series of sizes, each differing from the next by a similar factor. Each of these islands retains some areas representing the three major types of natural forest vegetation: lowland forest, montane forest, and mossy forest (or dipterocarp, lower montane, and upper montane rain forest; sensu Whitmore, 1 984). It is our purpose to assess the taxonomic status and summarize available ecological information for all mammal species known to occur on each island. Because many of these species are poorly known, we provide as much detailed information as is practical. Further analysis of distributional data is presented elsewhere (eg, Heaney et al, 1989; Heaney & Rickart, 1990; Heaney, 1991a). History of Investigations Although there are scattered early distributional records of mammals from Leyte (Hollister, 1912, 1913; Taylor, 1 934), the history of comprehensive collecting on these islands is limited. In 1963, D. P. Empress of Silliman University collected bats from within and around the Cathedral Cave complex near Inopacan, Hindang Municipality, Leyte (specimens in the Royal Ontario Museum). During the same decade, G. L. Alcasid and M. Celestino of the Philippine National Museum and D. S. Rabor of Silliman University made collections in the highlands of northern and central Leyte (specimens in the American Museum of Natural History and the Delaware Museum of Natural History). Most of these localities are mapped and briefly described by Parkes (1 973). Only small portions of these collections have been described in the literature (Heaney, 1985a; Musser, 1982ab; Peterson, 1981; Peterson & Fenton, 1970). Prior to our field studies (1981-1987), no surveys had been conducted on either Biliran or Maripipi. A preliminary listing of non-volant mammal species from all three islands appeared in Heaney (1986); however, this did not include the results from our extensive surveys of 1987. We have attempted to include information on all specimens now in museums. However, most of the information presented here is based on our own field surveys. Our work on Leyte was conducted along an elevational transect on the west-central portion of the island (figs. 1 2). A brief reconnaissance trip was made to this region from 23 to 28 May 1984 (voucher specimens in the University of Michigan Museum of Zoology ummz), but most work took place from 2 March to 13 April 1987 (specimens in the Philippine National Museum pnm, the Silliman University Museum su, the United States National Museum of Natural History usnm, the Visayas State College of Agriculture visca, and the Western Aus-Fig. 1 Map of Leyte Island, showing the relative positions of Biliran and Maripipi islands and the locations of Leyte collecting sites (as enumerated in text). A site of the elevational transect in the visca-mt. Pangasugan region (sites L1-L6) shown in Figure 2. Contour lines in meters. FIELDIANA: ZOOLOGY


Table 3. Extended. Prov: site L2 (2 usnm); site L4 (1 ummz); site L10 (5 amnh); site LI 9 (1 usnm). BILIRAN: site Bl (1 usnm). Order Chiroptera Family Pteropodidae — Fruit Bats Acerodon jubatus (Eschscholtz, 1831) The golden-capped flying fox is a Philippine endemic that is found throughout the country with the exception of the Palawan region. Gunther (1879) reported this species from southern Leyte. Specimens taken by Steere ( 1 890) on a small island midway in the channel between Leyte and Samar (site L20) were named Pteropus aurinuchalis (Elliot, 1896), but they are not now recognized as distinct. We did not encounter this species during our field work on Leyte, nor did we obtain any specimens on Biliran. We received reports of flying fox roosts on Biliran but did not locate any. On several occasions, however, we saw giant fruit bats flying at dusk over our campsites on Biliran or Table 4. Extended. RICKART ET AL: MAMMALS OF LEYTE, BILIRAN, AND MARIPIPI ISLANDS 23


Table 9. Extended. rain, during which time it groomed and licked off water droplets. The roost was not used on subsequent days. We collected lactating females in late April and early May. Males in our series are consistently larger than females in most cranial and external measurements (table 9). Specimens Examined— Total 20. MARIPIPI: site Ml (6 ummz); site M2 (6 ummz); site M3 (2 ummz); site M4 (5 usnm); site M10 (1 ummz). Other Records— LEYTE: site L7 (rom). Pteropus vampyrus lanensis Mearns, 1 905 any specimens, some of the bats we observed feeding in mabolo trees (Diospyros) on the campus of the Visayas State College of Agriculture (site LI) probably were this species. Local residents reported that large flying foxes were commonly seen at this site. There are no specimens from either Biliran or Maripipi. In the Leyte series, males are consistently larger than females in most cranial measurements (table 9). Specimens Examined— Total 14. LEYTE: Leyte Pro v: site L9 (10 dmnh); site L10 (2 amnh); specific locality unknown (2 fmnh). The giant flying fox is widespread in Southeast Asia and is locally common in primary forest throughout the Philippines. The species was first reported from Leyte by Steere (1890), and specimens have been recorded from Tambis and Patok (sites L9 and L10). Although we did not collect Rousettus amplexicaudatus amplexicaudatus (E. Geoffroy, 1810) The roussette fruit bat occurs from Thailand to the Solomon Islands and is found throughout the RICKART ET AL: MAMMALS OF LEYTE, BILIRAN, AND MARIPIPI ISLANDS 33


Table 12. Extended. 82 64 92 675 rump lengths 8-18 mm) were taken between 23 March and 27 April 1987. Males of this species are slightly larger than females in most cranial dimensions (table 1 2). Specimens from Leyte and Biliran are not distinguishable and are similar in size to specimens from Catanduanes (Heaney et al, 1 99 1 ; those from Negros are distinctly smaller (Ingle & Heaney, 1992). Specimens Examined— Total 88. LEYTE: Leyte Prov: site L2 (1 usnm); site L4 (2 su, 25 usnm); site L5 (6 usnm); site L6 (6 usnm); site LI 4 (5 dmnh). BILIRAN: site Bl (9 usnm); site B3 (2 pnm, 30 usnm, 2 wam). Rhinolophus rufus Eydoux & Gervais, 1836 This species is a Philippine endemic, with prior records from Bohol, Catanduanes, Luzon, Mindanao, and Mindoro (Heaney et al, 1991). We captured none, but in 1963 specimens were col-RICKART ET AL: MAMMALS OF LEYTE, BILIRAN, AND MARIPIPI ISLANDS 39


Table 14. Means (± SD) and ranges of selected measurements of adult vespertilionid bats (Murina, Myotis, Philetor, and Scotophilia) from Leyte, Biliran, and Maripipi islands. Note: Measurements other than weight are in millimeters. netted a single female on Biliran in partially logged primary forest at 700 m elevation (site B2; table 10). None were taken on Leyte or Maripipi. Measurements for the Biliran specimen are comparable to those for a male from Catanduanes (table 14; Heaney etal, 1991). Specimens Examined— Total 1. BILIRAN: site B2 (1 ummz). netted over a stream in the village of Viga (site M5). Two lactating females were captured in late April. Table 14 contains measurements for our specimens. Specimens Examined— Total 9. LEYTE: Leyte Prov: site L2 (2 usnm); site L4 (1 ummz). BILIRAN: site Bl (1 usnm); site B2 (3 ummz); site B3 (1 usnm). MARIPIPI: site M5 (1 usnm). Myotis muricola browni Taylor, 1934 This species is widespread in southern Asia and occurs throughout the Philippines (Hill, 1983; Koopman, 1989). On Leyte, we netted two specimens in disturbed lowland forest at 50 m and another in primary forest at 500 m (sites L2 and L4; table 10). Specimens from Biliran were taken in forest plantation at 50 m (site Bl), in partially logged primary lowland forest at 700 m (site B2), and in primary montane forest at 850 m (site B3; table 10). A single specimen from Maripipi was Philetor brachypterus (Temminck, 1 840) This bat occurs from Nepal to New Guinea. Previous Philippine records are from Mindanao and Negros. On Leyte, we netted two specimens in primary lowland forest at 500 m elevation (site L4; table 10). The species was not encountered on either Biliran or Maripipi. Measurements appear in Table 14. Specimens Examined— Total 2. LEYTE: Leyte Prov: site L4 (2 usnm). 42 FIELDIANA: ZOOLOGY


Table 14. Extended. Ear Forearm Weight (g) 47 38 74 98 38 15 36 It) Scotophilus kuhlii Leach, 1822 The widespread Asian house bat is abundant in most urban and agricultural areas throughout the Philippines. Our observations indicate that it is rare in forested habitats. On Leyte, it was abundant in the lowlands around the campus of the Visayas State College of Agriculture (site LI , where specimens were collected from a roost of several hundred in the attic of a large building. Five others were netted along a stream in secondary forest at 50 m elevation at the base of Mt. Pangasugan (site L2; table 10). On Biliran, we netted house bats and collected them from roosts in buildings in the town of Naval (site B5). Here they were also seen roosting under the dead fronds of an ornamental fan palm, Livistona rotundifolia. Some of the fronds may have been modified to form "tents; S. kuhlii is the only vespertilionid known to exhibit this behavior (Rickart et al, 1989b). On Maripipi, one specimen was netted at 400 m elevation on a ridge between areas of secondary lowland forest (site Ml; table 10), and six others were netted over a stream in the village of Viga (site M5). Females collected during April all contained two embryos. Pregnancies among a series of 14 females taken from a single colony on 27 April were relatively asynchronous (crown-rump lengths 7-2 1 mm). Specimens from Leyte have slightly larger crania than do those from Biliran (table 14). Specimens Examined— Total 89. LEYTE: Leyte Prov: site LI (13 usnm); site L2 (5 usnm); site LI 5 (15 amnh); site LI 6 (22 fmnh). BILIRAN: site B5 (2 pnm, 1 su, 9 ummz, 14 usnm, 1 visca). MARIPIPI: site Ml (1 ummz); site M5 (6 usnm). Other Records— LEYTE: site LI 7 (rom). Family Molossidae — Free-tailed Bats Chaerephon plicata (Buchanon, 1 800) This species of free-tailed bat is found from India to Southeast Asia. Chaerephon, formerly a sub-RICKART ET AL: MAMMALS OF LEYTE, BILIRAN, AND MARIPIPI ISLANDS 43


Table 19. Extended. Specimens Examined— Total 54. LEYTE: Leyte Prov: site LI (2 usnm); site L2 (1 usnm); site LI 5 (4 amnh); site L9 (10 dmnh); site L10 (7 amnh); site LI 3 (1 dmnh); site LI 6 (4 fmnh). BILIRAN: site Bl (4 usnm); site B2 (3 ummz); site B3 (5 usnm); site B4 (3 ummz). MARIPIPI: site Ml (4 ummz); site M4 (5 usnm); site M10 (1 ummz). Order Carnivora Family Viverridae— Civets te, we trapped one animal in primary lowland forest at 320 m elevation (site L3) and shot a second individual from a tree in an area of mixed agriculture and second growth at 50 m elevation (site L2). We received reports of civets on both Biliran and Maripipi but obtained no specimens. The descriptions given by most informants conformed to that of the palm civet. On both islands we also found civet scats that most closely resembled those of Paradoxurus (ie, composed primarily of seeds; Heideman et al, 1987). Specimens Examined— Total 2. LEYTE: Leyte Prov: site L2 (1 usnm); site L3 (1 usnm). Paradoxurus hermaphroditus philippinensis Jourdan, 1837 Viverra tangalunga Gray, 1832 The palm civet occurs from India to Southeast Asia and is widespread in the Philippines. On Ley-The Malay civet (or tangalung) is found from peninsular Malaysia to Sulawesi. Although it is RICKART ET AL: MAMMALS OF LEYTE, BILIRAN, AND MARIPIPI ISLANDS 51


Table 20. Continued. Artiodactyla Cervus mariannus Sus barbatus Native species Non-volant species Large fruit bats Small fruit bats Microchiroptera All natives Introduced species All species Distributions (after Heaney et al. 1987): LEYTE restricted to Leyte; MIND restricted to Mindanao faunal region; PHIL widespread in oceanic Philippines; WIDE occurring outside of oceanic Philippines, t Introduced commensal species. typically have low population densities and thus require larger areas to sustain viable populations. We believe it likely that the absence of pigs and deer on Maripipi stems from natural extinction. However, it is certainly possible that both species may have become extinct through relatively recent hunting pressures. Small mammals with restricted elevational ranges also appear to be susceptible to extinction on small islands, presumably because of narrower ecological tolerances. On Leyte, Batomys salomonseni was restricted to high-elevation forest, where it was the most frequently trapped species (table 1). However, it was rare in similar habitat on Biliran and was absent from Maripipi. The distribution of Apomys littoralis is suggestive of similar limitations. The situation for the widespread Philippine native rat, Rattus everetti, provides an interesting contrast. In surveys on Leyte (this study, table 1 ) and Luzon (Rickart et al, 1 99 1 , this species was common in habitats ranging from lowland dipterocarp forest to high-elevation mossy forest. Perhaps because of this broad ecological tolerance, it has survived on Maripipi, where more specialized species have not. Ecological factors may also explain the distributional limitations of small fruit bats among these islands. Of the species that were common on Leyte and Biliran, Haplonycterisfischeri and Ptenochirus minor were conspicuous in that they were almost never encountered outside of forested habitats. This inability to tolerate disturbance may be associated with their apparent absence from Maripipi. Netting and Trapping Success Our field work on Leyte served as a basis for formulating several general predictions concerning patterns of variation in species richness and abundance of small mammals in Southeast Asia (Heaney et al, 1 989). Subsequent studies generally have supported these predictions (Heaney & Rickart, 1 990; Heaney et al, 1 99 1 ; Rickart et al, 1 99 1 , and we believe that data from Biliran and Maripipi can be used to further test their generality. The relevant predictions from Heaney et al. (1989) derived from our studies on Leyte and Negros are (1) in forested habitats, species richness and relative abundance of fruit bats should be highest in the lowlands and decline with increasing elevation, (2) species richness of fruit bats should be lower in agricultural areas, where geographically widespread species should predominate, (3) non-volant mammals should not exhibit highest species richness in lowland forest, (4) abundance of non-volant mammals should increase with elevation, and (5) endemic species of fruit bats and non-volant mammals should occur in primary forest, whereas non-endemics should predominate in disturbed habitats. Data on species richness of small fruit bats on Biliran and Maripipi (table 5) generally support predictions 1 and 2. Species richness was highest at the lowest elevation forested sites (sites B2 and Ml) and declined with elevation to reach lowest values in mossy forest. On both islands there was also an indication of a decline in relative abunrickart ET AL: MAMMALS OF LEYTE, BILIRAN, AND MARIPIPI ISLANDS 57