Papers by Lawrence R Heaney
Biological Reviews (Cambridge Philosophical Society), 2016
The general dynamic model of oceanic island biogeography (GDM) has added a new dimension to theor... more The general dynamic model of oceanic island biogeography (GDM) has added a new dimension to theoretical island biogeography in recognizing that geological processes are key drivers of the evolutionary processes of diversification and extinction within remote islands. It provides a dynamic and essentially non-equilibrium framework generating novel predictions for emergent diversity properties of oceanic islands and archipelagos. Its publication in 2008 coincided with, and spurred on, renewed attention to the dynamics of remote islands. We review progress, both in testing the GDM's predictions and in developing and enhancing ecological–evolutionary understanding of oceanic island systems through the lens of the GDM. In particular, we focus on four main themes: (i) macroecological tests using a space-for-time rationale; (ii) extensions of theory to islands following different patterns of ontogeny; (iii) the implications of GDM dynamics for lineage diversification and trait evolution; and (iv) the potential for downscaling GDM dynamics to local-scale ecological patterns and processes within islands. We also consider the implications of the GDM for understanding patterns of non-native species diversity. We demonstrate the vitality of the field of island biogeography by identifying a range of potentially productive lines for future research.
National Museum of the Philippines: Journal of Natural History , 2013
ABStrAct We conducted a survey of non-volant small mammals in 2006-2008 in four areas of the Bico... more ABStrAct We conducted a survey of non-volant small mammals in 2006-2008 in four areas of the Bicol Peninsula: Mt. Labo (peak 1544 m), Mt. Malinao (peak 1548 m), Saddle Peak (peak 1003 m), and Caramoan National Park (475 m). In 11,227 trap-nights we documented nine species, of which six were native and three were introduced. The native species consisted of one shrew and five rodents; the exotic species included one shrew and two rodents. Species diversity was comparatively low overall, with each mountain supporting from three to four species. None of the four species previously documented on Mt. Isarog (Archboldomys luzon-ensis, Batomys sp., Chrotomys gonzalesi, and Rhynchomys isarogensis) were present in these newly surveyed areas; the Chrotomys from Saddle Peak and Rhynchomys from Mt. Labo are of uncertain identity. We did not record any species of large Apomys (subgenus Megapomys), which are abundant in central and northern Luzon, indicating that these forest mice are absent on the Bicol Peninsula. We captured the exotic rodents only in heavily disturbed forest and subsistence farms in the lowlands; we found the introduced Asian house shrew, Suncus murinus, only in montane forest on Mt. Labo, ca. 1335-1413 m. Relative abundance of the native species was low overall, ranging from 0.32 to 3.31 individuals/100 trap-nights. The presence of the two possibly new species of Chrotomys and Rhynchomys, in addition to the four species endemic to Mt. Isarog, highlight the uniqueness of the Bicol mammal fauna. We recommend that Saddle Peak be designated and managed as a protected area similar to the other areas we surveyed for its importance as a watershed for the municipalities of Camarines Sur surrounding it and as habitat of endemic mammals.
National Museum of the Philippines: Journal of Natural History , 2013
An intensive survey of the small non-volant mammals along the full elevational gradient of Mt. Ba... more An intensive survey of the small non-volant mammals along the full elevational gradient of Mt. Banahaw -San Cristobal National Park in 1995-2005, and 2012, using standardized procedures, produced records of ten native and three exotic small mammals. Four species discovered during our survey appear to be endemic to the park (Apomys banahao, A. magnus, Musseromys gulantang, and Rhynchomys banahao), a remarkable number for such a small area; the other native species are more widespread. Species richness in forest on Mt. Banahaw (peak = 2169 m) increased from five at 620 m to seven at 1465 m, and declined to three at 2030 m. Mixed agricultural areas and exotic grassland with shrubby second growth supported up to three exotic species and up to three native species. Exotic species were apparently absent from secondary or primary forest. Relative abundance of native species was significantly correlated with increasing elevation, with abundance about five times greater in montane and mossy forest than in the lowlands. Trap success using live earthworms as bait increased significantly with increasing elevation, and small mammals showed an overall preference for earthworm bait, but a few were arboreal and a few preferred coconut bait. Most species were significantly more active on the ground than in trees, and most were significantly more active during the night than during daylight. Mt. Banahaw -San Cristobal National Park is a premier example of a highly localized center of mammalian endemism; along with its important role as a religious and recreational center and crucial function as a watershed, it clearly deserves continued and active protection.
National Museum of the Philippines: Journal of Natural History , 2013
Field surveys of the mammal fauna of Mt. Natib, Bataan Province, conducted in 1996 and 2005 docum... more Field surveys of the mammal fauna of Mt. Natib, Bataan Province, conducted in 1996 and 2005 documented a total of 20 species including nine bats, five native non-volant small mammals, two non-native small mammals, and four native large mammals. Several additional species of bats and two additional large mammals that are known to occur elsewhere in southwestern Luzon also probably occur on Mt. Natib. One native species, Apomys zambalensis, is endemic to Mt. Natib and the nearby Zambales Mountains and was the most abundant small mammal present. Survey results for non-volant mammals were consistent with predicted diversity of native species for a mountain of relatively low elevation, and strengthened a general pattern of a positive correlation between local species richness and elevation on Luzon. The occurrence of non-native species was strongly associated with highly disturbed habitats. Native species occurred across a broad range of disturbance conditions and were numerically dominant over non-natives even in the most disturbed situations. Results suggest that the Natib mammal fauna is highly tolerant of anthropogenic disturbance, perhaps reflecting a legacy of coping with periods of severe volcanic disturbance.
National Museum of the Philippines: Journal of Natural History , 2013
Mammalia, 2016
Faunas of old oceanic islands often have extremely high levels of endemism and are considered hig... more Faunas of old oceanic islands often have extremely high levels of endemism and are considered highly susceptible to anthropogenic disruption. We surveyed the richly endemic small mammal fauna on Mt. Amuyao in the Central Cordillera of northern Luzon Island, Philippines. We tested hypotheses regarding elevational patterns of species richness and community composition, community response to habitat disturbance, and interactions of native and non-native mammals. Our study revealed greater species richness and faunal hetero-geneity within the Central Cordillera than previously suspected. We documented 15 native species (14 rodents and 1 insectivore), and two species of non-native rodents. All of the native species are endemic to the Philippines, eight being restricted to the Cordillera. Twelve of the 14 native rodents belong to two ancient endemic clades, indicating that most of the regional diversity is the product of in situ speciation. Native mammal assemblages are ecologically diverse, and include species with varied trophic habits, activity patterns, and climbing ability. Some native species are restricted to relatively pristine habitat, whereas others are highly tolerant of disturbance. Non-native species are restricted to highly disturbed habitats and apparently are displaced by natives where habitat has regenerated from past disturbance.
National Museum of the Philippines: Journal of Natural History, 2013
National Museum of the Philippines: Journal of Natural History, 2013
Frontiers of Biogeography (Int. Biogeo. Soc.), 2013
Models of biogeographic processes can both enhance and inhibit our ability to ask questions that ... more Models of biogeographic processes can both enhance and inhibit our ability to ask questions that guide our understanding of patterns and processes. The two 'traditional' models of island biogeography, the Equilibrium Model and the Vicariance Model, raise important and insightful questions about relevant processes, but both fail to raise many crucial questions. An example involving the non-volant mammals of the Philippine archipelago shows that both models highlight some, but not all, relevant patterns and processes. The more recently proposed General Dynamic Model successfully combines many of the positive aspects of the two traditional models, but leaves some important questions unasked. We pose a number of questions here that may help guide further development of models of island biogeography. perspective 250 frontiers of biogeography 5.4, 2013 -
Diversity and Distributions , 2011
Aim Island faunas, particularly those with high levels of endemism, usually are considered especi... more Aim Island faunas, particularly those with high levels of endemism, usually are considered especially susceptible to disruption from habitat disturbance and invasive alien species. We tested this general hypothesis by examining the distribution of small mammals along gradients of anthropogenic habitat disturbance in northern Luzon Island, an area with a very high level of mammalian endemism.
New Perspectives on the Long-Term Biogeographic Dynamics and Conservation of Philippine Fruit Bats, 2009
a Faunal regions (fig. 2.2) are A = Greater Luzon; B = Greater Mindanao; C = Greater Negros-Panay... more a Faunal regions (fig. 2.2) are A = Greater Luzon; B = Greater Mindanao; C = Greater Negros-Panay; D = Greater Palawan; E = Mindoro; F = Sibuyan; G = Babuyan/Batanes. b Forest habitat types are L = lowland; M = montane; H = highland mossy.
J. Int. Biogeo. Soc. , 2009
Islands that have been isolated since their formation or for very long periods of time often have... more Islands that have been isolated since their formation or for very long periods of time often have land mammal faunas that are made up largely or entirely of endemic species, and often these species are members of speciesrich endemic clades. Though members of these endemic clades are each other's closest relatives, they typically show highly diverse body size, morphology, behavior, and ecology. These constitute classic cases of adaptive radiation, in which local speciation has produced spectacular diversity.
The Mammals and Birds of Camiguin Island, Philippines, a Distinctive Center of Biodiversity , 2006
Fifty-five species of resident breeding and two species of migratory land birds have been documen... more Fifty-five species of resident breeding and two species of migratory land birds have been documented on Camiguin Island, including one bittern, one eagle, one junglefowl, two rails, eight doves and pigeons, one parrot, three cuckoos, one owl, three swiftlets, one dollarbird, two kingfishers, one bee-eater, one hornbill, one pitta, one triller, two bulbuls, one crow, four thrushes, three warblers, six flycatchers, one pipit, one wood swallow, two starlings, three sunbirds, two flowerpeckers, two white-eyes, and two munias. At least seven species reported here are first records for Camiguin. Ten species are widespread Philippine endemics, two are near-endemics, and one (Loriculus camiguinensis, described in this volume) is endemic to Camiguin. Additionally, four endemic subspecies are recognized from Camiguin, Ixos everetti catarmanensis, Hypothymis azurea catarmanensis, Diceum trigonostigma isidroi, and Zosterops nigrorum catarmanensis. While this list is not comprehensive, the presence of 57 species demonstrates that many species were able to cross a narrow but permanent sea channel, and the presence of four endemic subspecies and one endemic species indicates that some genetic differentiation has resulted.
The Mammals and Birds of Camiguin Island, Philippines, a Distinctive Center of Biodiversity , 2006
Biodiversity surveys in the 1960s and 1990s on Camiguin Island, a geologically young, volcanicall... more Biodiversity surveys in the 1960s and 1990s on Camiguin Island, a geologically young, volcanically active oceanic island surrounded by deep water, have demonstrated the presence of 24 species of land mammals. Three species (one insectivore and two rodents) are not native to the Philippines, but all others (one insectivore, 12 bats, one monkey, four rodents, two small carnivores, and one ungulate) are indigenous. Among those captured in the 1990s were two previously unknown species of murid rodents in the genera Apomys and Bullimus that are endemic to Camiguin. The discovery of two new species on such a small island (ca. 265 km 2) is remarkable; Camiguin is currently the smallest island in the Philippines known to have unique species of mammals. Total species richness of nonvolant mammals on Camiguin is low, but relative to islands that were once part of Pleistocene Greater Mindanao, Camiguin is not depauperate. However, its fauna is not ecologically balanced in the same way as the faunas of the islands that were part of Greater Mindanao: ground-living shrews (Crocidura) and rodents (Apomys, Bullimus, Crunomys, and Rattus) from lowland forest, along with some large mammals (Macaca, Paradoxurus, and Sus) are well represented on Camiguin, but all the arboreal small mammals that characterize lowland forest on Mindanao (Sundasciurus, Exilisciurus, Cynocephalus, and Tarsius), ground-living small mammals from montane habitats (Urogale, Podogymnura, Batomys, Limnomys, and Tarsomys), and one large mammal (Cervus) are absent. Additionally, at least two genera of fruit bats (Haplonycteris and Megaerops) that are fairly common in lowland rain forests on Mindanao are absent on Camiguin. The presence of some nonvolant mammals demonstrates that dispersal across the deep but narrow intervening channel takes place, but the presence of two species endemic to Camiguin and the absence of other species that are present on nearby Mindanao implies that dispersal probably is rare. The Asian house shrew (Suncus murinus) was remarkably abundant in primary forest at high elevation; this species has also been found to be abundant in montane primary forest on Negros Island, which also has low total species richness. Species richness of small nonflying mammals was greatest at fairly high elevation.
The Mammals and Birds of Camiguin Island, Philippines, a Distinctive Center of Biodiversity , 2006
An inventory of the mammals of Camiguin Island conducted in 1994 and 1995 documented the presence... more An inventory of the mammals of Camiguin Island conducted in 1994 and 1995 documented the presence of a previously unknown species of Philippine forest mouse of the endemic Philippine genus Apomys, which is here named and described. Based on molecular data, the new species is most closely related to two species (A. hylocoetes and A. insignis) from Mindanao Island and to an unnamed species from Leyte, Biliran, and Bohol islands. The new species is diagnosed in comparison to its three closest relatives on the basis of slightly browner and less russet fur, slightly greater size overall, a moderately long and broad hind foot with small plantar pads, large tail scales, slightly narrower zygomatic width and mastoid breadth, deep rostrum of moderate length, a long orbit and braincase, narrow palate, large incisive foramina, short distance from the posterior edge of the incisive foramina to the anterior edge of the first upper molar, bony palate that extends well to the posterior of the posterior edge of the last upper molar, bullae that are more strongly oriented toward the cranial midline axis, third upper molar without a conspicuous anterolabial cusp, and a number of more subtle features. It is one of two species of mammals now known to be endemic to Camiguin, the other being Bullimus gamay (Rickart et al., 2002). Both are common in rain forest on Camiguin Island at upper elevations. The presence of two endemic mammals on this small (265 km 2) island is remarkable; there are no smaller islands in the Philippines known to support endemic mammal species.
The Mammals and Birds of Camiguin Island, Philippines, a Distinctive Center of Biodiversity , 2006
Camiguin Island, with an area of ca. 265 km 2 and maximum elevation of ca. 1620 m, lies about 10 ... more Camiguin Island, with an area of ca. 265 km 2 and maximum elevation of ca. 1620 m, lies about 10 km north of Mindanao but is isolated from Mindanao by a deep (385 m) channel. It originated from volcanic activity as a dryland island not earlier than 1 million years ago, but most growth of the island has occurred within the past 340,000 years. Current landforms are dominated by large, scenic volcanic peaks, several of which are active. Lowland rain forest originally occurred up to about 1100 m elevation, with montane rain forest from 1100 m to about 1350 m and mossy forest from 1350 m to the peaks. By the mid-1990s, deforestation had removed most natural vegetation below about 600 m, with degree of disturbance to forest decreasing with elevation and ending at about 1250 m. The climate is tropical, with rainfall of 2-3 m per year in the lowlands and probably about 7.5 m near the peaks. Mammal and/or bird specimens are available from 18 sites from the 1960s and 1990s; these sites are here located and described to the extent possible. Given the presence of two endemic species of mammals (one described in this volume), one endemic bird (described in this volume), and previously described endemic plants and a frog, Camiguin is one of the smallest but most distinctive centers of biodiversity in the Philippines and should be a priority site for conservation. The remaining forest on Camiguin is essential habitat for these unique species, but it is also essential for watershed protection and control of floods and landslides, and it contributes significantly to the tourism trade that provides substantial income on the island. Deforestation for logging and agriculture and overhunting are current threats. A protected area on the island should include the full range of original habitat diversity, which would encompass both the existing high-quality forest at upper elevations and also significant tracts of disturbed but natural lowland forest, especially along rivers and streams, that should be allowed to regenerate in the future.
The Mammals and Birds of Camiguin Island, Philippines, a Distinctive Center of Biodiversity , 2006
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Papers by Lawrence R Heaney