Article Interactions between riparian vegetation and fluvial processes within tropical Southeast Asia: Synthesis and future directions for research Progress in Physical Geography 2014, Vol. 38(6) 716–733 ª The Author(s) 2014 Reprints and permission: sagepub.co.uk/journalsPermissions.nav DOI: 10.1177/0309133314548745 ppg.sagepub.com Helen L. Moggridge University of Sheffield, UK David L. Higgitt University of Nottingham Ningbo China, China Abstract Research on the interaction between vegetation and hydrological and geomorphological processes has made a significant advancement in river and floodplain research. While this work is strongly dominated, both conceptually and empirically, from studies in northern temperate systems, it has been instrumental in shaping understanding at a global scale. There are, however, regions of the world such as tropical Southeast Asia which have received relatively little research attention, but could offer an important contribution to knowledge in this field. In a first step to address this issue, this paper synthesizes current research on vegetation and geomorphology and hydrology within tropical Southeast Asia, to consider the applicability of current (temperate-based) models and the potential contribution that processes within these systems could make to global understanding. While research within the region is sparse, observations suggest a reciprocal relationship between vegetation and fluvial processes. While there are some synergies with temperate systems, processes within this region also present interesting differences which could, with further investigation, advance current understanding of these processes globally and expand and enhance current concepts. The paper concludes with the identification of the pertinent research questions for the field within the region. Keywords fluvial geomorphology, hydrology, large wood, Southeast Asia, tropical, vegetation I Background and introduction The reciprocal role between vegetation and physical processes in rivers and their wetlands has received considerable research interest in recent decades, both conceptually (e.g. Corenblit et al., 2007; Fisher et al., 2007) and empirically (e.g. Bertoldi et al., 2011). Studies have shown that hydrological and geomorphological processes can influence vegetation composition (Bendix and Hupp, 2000), by creating a mosaic of heterogeneous habitats (which vary in disturbance frequency and intensity and nutrient and moisture availability) and through facilitating Corresponding author: Helen L. Moggridge, Department of Geography, Winter Street, University of Sheffield, Sheffield S10 2TN, UK. Email: [email protected] Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 Moggridge and Higgitt 717 the dispersal, establishment and growth of vegetation (Greet et al., 2011; Steiger et al., 2005). The active physical role that established vegetation can have in subsequently modifying these physical processes has also been well studied; vegetation modifies water flows and sediment dynamics (e.g. Tabacchi et al., 2000), causing rivers to develop new landforms (e.g. Bertoldi et al., 2011), alter meander migration rates (e.g. Perucca et al., 2007), influence braiding patterns (Coulthard, 2005; Tal et al., 2004) and create anastomosing planforms (Tooth and Nanson, 1999, 2000). This reciprocal relationship is comprehensively reviewed in temperate river systems by Corenblit et al. (2009a) and Gurnell et al. (2012). The foundation for much of this research was the notable and widely cited Flood Pulse Concept (Junk et al., 1989), which introduced the idea that lateral connectivity within fluvial systems, through periodic inundation of floodplains, was ecologically significant. This was based on field data for fish communities in large rivers in the Neotropics and North America and has underpinned subsequent research on physical and ecological linkages within river systems worldwide (Junk and Wantzen, 2004). The concept has since been advanced in tropical rivers; fish ecology has been explicitly linked with particular aspects of the flood pulse, such as the timing, continuity (i.e. absence of any disruption from floods), smoothness (i.e. steadiness of the rise and fall), rapidity of change amplitude and duration of the flood pulse (see Welcomme and Halls, 2004, and references therein). Research has also broadened to consider riparian vegetation; several studies on the Parana and Amazon rivers have demonstrated that the flood pulse interacts with geomorphological processes to create a heterogenous mosaic of different landforms and wetland types, which all support distinct vegetation communities and contribute to the diversity of these systems (Hamilton et al., 2007; Kalliola et al., 1991; King, 2003; Lamotte, 1990; Marchetti et al., 2013; Mertes et al., 1995; Stevaux et al., 2013). Research from tropical Africa shows similar relationships (Hughes, 1990; Medley, 1992, and references therein), although few studies exist. The tropical origin of the Flood Pulse Concept is, however, rather anomalous in river research. While there have been notable advances in research within the tropics (and across all ecoregions), northern temperate systems have received a particular focus. Most of the key models that precede (e.g. River Continuum Concept, Vannote et al., 1980) or follow on from the Flood Pulse Concept (e.g. Shifting Habitat Mosaic, Stanford, 1998; Flow Pulse Concept, Tockner et al., 2000; River Ecosystem Synthesis, Thorp et al., 2006) are heavily based on research within the temperate zone and these models are being used to guide research questions and management approaches in rivers worldwide (Boulton et al., 2008). The strong geographical foci of this research constrains understanding to a limited number of climates, flow regimes and vegetation types, which ultimately limits the depth of understanding of these processes and leaves many systems under-researched. Key examples are the Southeast Asian monsoonal river and floodplain systems. These systems are characterized by exceptionally high biodiversity (e.g. Keogh et al., 1999) and deliver important ecosystem services (Jusoff and Majid, 1990), but are acknowledged as being some of the least studied biotopes in the region (Anderson, 1983; Dudgeon, 2000a). Further, the research that is conducted in this region rarely captures a global audience; ‘[studies] dealing with tropical rivers seem generally to be perceived as interesting regional studies first and contributions to limnology second; studies of north-temperate waters are viewed in the opposite way’ (Dudgeon, 2000a: 258). This lack of research is also confounded by the extensive degradation and decline of these systems (Dudgeon, 1992, 2000b). Thus, there is an urgent and important need for future research to consider these Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 718 Progress in Physical Geography 38(6) 120°E 100°E 140°E Mek utr ong Brahmap a C h i n a Irr awa ddy Bangladesh Burma 20°N 20°N Thailand Cha o Phr ay a Loas Vietnam Philippines Cambodia Pacific Ocean M a l a y s i a Equator Borneo Sumatra Papau New Guinea I n d o n e s i a Fl y Java Indian Ocean 0 1000 km 20°S 100°E Australia 120°E 140°E 20°S Figure 1. Map of the study area, Southeast Asia and the surrounding tropical monsoon regions. The largest rivers of the region are shown. systems, to investigate the applicability of existing models, to contribute to global understanding of vegetation and fluvial processes and to inform effective management. This paper will synthesize existing studies within tropical Asia to develop a basic conceptual understanding of the physical role of vegetation in rivers and their wetlands. The focus area of the study (shown in Figure 1) are rivers within monsoonal Southeast Asia, with consideration, where appropriate, of tropical rivers extending to India, Papua New Guinea and tropical northern Australia. Relevant studies from the wider tropical latitude of 23 N and 23 S are also used to support discussion. The paper will present an initial comparative evaluation of the synergies and contrasts between models derived from the temperate zone and observation from tropical rivers and identify the principal research directions for future study. While ecological comparisons have been made between temperate and tropical streams previously (see Boulton et al., 2008, for an overview) and the geomorphology of tropical rivers has been reviewed (see Latrubesse et al., 2005, for a global review of tropical systems and Gupta, 2005, for a regional overview), there is little available information on rivers within this region. Further, vegetation within rivers and its interaction with fluvial processes have not been previously considered in Southeast Asia, with existing work on tropical rivers being almost entirely confined to the Neotropics. Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 Moggridge and Higgitt 719 II Southeast Asian rivers and floodplains A principal distinction between temperate and tropical rivers is the flow regime, which is critical to determining vegetation distribution, survival and growth (Greet et al., 2011). Southeast Asia exhibits a wet tropical climate, with annual rainfall typically exceeding 2500 mm. This drives the flow regime, as all the rivers within the region are rain fed (Gupta, 2005). Seasonality within the region is driven by two opposite patterns of monsoon winds and so can vary from a long, wet season (9–10 months duration) and a shorter dry season, or two monsoon seasons (October–March and April–August) (Page et al., 2006). Close to the equator, seasonality is less pronounced, but still evident from the variation in the Intertropical Convergence Zone. This heterogeneity in seasonality is reflected in the flow regimes of the river catchments. For example, small equatorial catchments often exhibit little seasonality, with a flashy variation in discharge, mostly attributable to individual high-magnitude, shortduration storm events (Spencer et al., 1990). In contrast, many of the larger systems are profoundly influenced by the monsoon, which creates a characteristic pattern of well-defined wet and dry seasons, usually with the flood peak in August–September (Dettinger and Diaz, 2000) although with variation in timing across the region (Dudgeon, 1992). The intensity of the monsoon is linked with the El Niño-Southern Oscillation phenomenon, which drives more interannual variability in flood magnitude in this region than is typically associated with tropical rivers (Boulton et al., 2008; Puckridge et al., 1998). This is also confounded by the influence of typhoons and tropical cyclones (Boulton et al., 2008), whose seasonal arrival coincides with the high flow period and can cause large-scale flooding. A defining characteristic of catchments within this region is the large variation in size; five of the world’s longest rivers are in tropical Asia (Mekong, Indus, Brahmaputra, Ganges and Irrawaddy). The largest river within the region is the Mekong River, which drains 795,000km3 and has a mean discharge of 15,000 m3s-1 at the mouth (Gupta et al., 2002), classifying it as a ‘mega-river’ (Latrubesse, 2008). The monsoon rainfall drives the flow regime of these major river systems, but the characteristics and behaviour of these rivers are also fundamentally influenced by tectonics and Quaternary sea-level changes (Gupta, 2005). The region is characterized by rapid weathering, driven by heavy rainfall, a steep topography, erodible rocks and high levels of tectonic activity. This generates high sediment loads, particularly in the rivers draining large mountainous areas, such as the Brahmaputra, Mekong and Irrawaddy, and rivers draining the highelevation oceanic islands, including the Fly and Sepik in Papua New Guinea and small rivers draining Java and Borneo (Latrubesse et al., 2005). Sediment transport is tightly coupled to the rainfall regime and thus strongly pulsed with the heavy rainfall events (often tropical cyclones) that characterize the region. Sediment erosion, transport and deposition (and subsequently all channel-defining processes) exhibit strong seasonality associated with the wet season (Gupta, 2011); it has been suggested that over 50% of the sediment eroded within a tropical catchment can be transported in just a few days in the year (Douglas, 1993). An initial classification of river types in tropical South East Asia is proposed in Table 1, based on a very limited collection of morphological descriptions of rivers within the region. The classification distinguishes between confined, bedrock-controlled rivers and alluvial rivers with extensive floodplains. Confined systems include small, steep rivers draining mountainous and volcanic areas, and confined sections of larger rivers (e.g. Figure 2), which are common within the region. Three major types of alluvial rivers are identified: meandering, Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 720 Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 Floodplain morphology Characteristic wetlands and riparian vegetation assemblages B: Lowland, alluvial rivers with extensive floodplains Seasonally inundated wetlands on scroll B1: Single channel, meandering Floodplain morphology comprising scroll complexes, abandoned bars and alluvial plains. channels, oxbow lakes, alluvial plains Lower elevations characterized by of minor tributaries and permanently inundated wetlands, such as backswamps and blocked backswamps. valley swamps, and open-water habitats, such as oxbow lakes. Point and channel bars and islands occur Pioneer vegetation may colonize bar B2: Braided planform, often within and adjacent to main channel. tops. due to a high sediment supply from draining Natural levees may occur along Seasonally inundated wetlands on levee mountainous reaches channel banks. tops. Permanently inundated wetlands, such as swamps and shallow lakes, occur between levees and floodplain. Periodically inundated vegetation on B3: Anabranching planforms, Large bars and islands within the island tops. characteristic of the channel, often with well-developed Seasonally inundated wetlands on levee lower reaches of the vegetation. Floodplains charactervery large river systems ized by scroll bars, crevasse splays tops, scroll bars. and levees (based on studies from Permanently inundated wetlands at lower elevations, such as large Neotropical, anabranching backswamps and open-water systems). habitats, such as shallow lakes. A: Confined rivers with little or no floodplain A1: Steep upland, headwater Rivers are confined by steep banks, with No riparian wetlands. Riparian streams no floodplain, especially in upstream vegetation confined to a narrow reaches. Lateral bars may form margin along the main channel. further downstream. Possible vegetation colonization of A2: Narrow, incised rivers No floodplain, rivers confined and often in-channel bars and islands. draining volcanic areas deeply incised. Prone to significant modification from episodic sedimentation from volcanic activity. A3: Bedrock channels in Little or no floodplain development, confined sections of with channels flowing through large rivers gorges. In wider sections, rock cored in-channel islands may be present (e.g. Mekong) and bar deposits may overlay bedrock (e.g. Irrawaddy). Category description Lower reaches of Irrawaddy, Ganges and Mekong at Laos-Cambodia border (‘4000 Islands’). Gupta and Liew (2007); Jain et al. (2012); Marchetti et al. (2013); Stevaux et al. (2013) Sections of the Brahmaputra in Assam, Dietrich et al. (1999); Sarma northwest India. (2005) Wider, mountainous reaches in upper sections of the Fly River, Papua New Guinea. Lower reaches of Fly River, Papua New Deitrich et al., (1999); Gupta and Liew (2007); Gupta et al. Guinea. Mid- to lower reaches of (2002); Pickup (1984); Pickup Chao Phraya, Thailand. Reaches of and Marshall (2009) upper Mekong (downstream of Vientiane, Lao PDR). Gupta (2005); Gupta and Liew (2007); Gupta et al. (2002); Stamp (1940) Gupta (2005) Rivers draining volcanic areas of Sumatra and Java. Sections of Irrawaddy and upper Mekong (downstream of Savannakhet, Lao PDR). Upper reaches of Chao Phraya, Thailand. Gupta (2005) References Headwaters of the Pahang, Malaysia. Examples Table 1. Classification of the major river types within Southeast Asia, with their characteristic wetlands and vegetation assemblages, with key examples. Moggridge and Higgitt 721 Figure 2. Rock-cored in-channel island of the Mekong River, upstream of Chiang Sean, Thailand (A3). Fine sediment deposited on and between the rock outcrops creates suitable habitat for colonization of vegetation. braided and anabranching. The classification is not intended to be exhaustive, but merely serves as a framework for discussion. Of particular note is that the morphology of these rivers can vary within systems as much as between them—a characteristic of tropical rivers worldwide (Latrubesse et al., 2005)—with many larger rivers alternating between bedrockconstrained channels and alluvial channels (Gupta, 2005) and thus appearing as several river types in Table 1. For example, the 4880 km course of the Mekong changes from constrained narrow rocky valleys to wide alluvial basins with meandering planforms (Gupta, 2011), with a complex 4000-island archipelago at the Lao PDR/Cambodia border which creates an anabranching planform (Latrubesse et al., 2005). The smaller Fly River in Papua New Guinea (catchment area 75,000 km3) exhibits three distinct morphological zones along its course (Pickup, 1984): a gorge section in the mountainous upper reaches, which can be up to 1000 m deep and has an active hillslope sediment supply, causing poorly developed braiding in the wider sections; a valley section, characterized by a braided channel, interspersed with vegetated islands; and a meandering section in the lowland alluvial plains, characterized by a single channel with a floodplain morphology derived from past channel positions (Dietrich et al., 1999). In certain sections, rivers may traverse river types too; for example, some meandering sections of the Mekong exhibit bar-braided planforms at low flows (Gupta and Liew, 2007) and the Brahmaputra has braided, anabranching reaches (Sarma, 2005). Within the active zone of all of these rivers, flow and sediment dynamics interact to create a diverse mosaic of vegetation communities, particularly in the low-gradient rivers with well-developed floodplains. There is little information on the riparian vegetation of the volcanic and steep, upland rivers (types A1 and A2), although a description of upland streams in Kalimantan (Indonesian Borneo) suggests a diverse riparian fringe vegetation composed of aroids and leguminous trees (MacKinnon et al., 1996). No published information was available on vegetation within the confined sections of larger rivers (A3), although observation of a rock-confined section of the Mekong showed vegetation establishment on in-channel islands (see Figure 2).The floodplains of the lowland, alluvial rivers (types B1–B3) are naturally dominated by rainforest vegetation, which is strongly associated with the flow regime of the catchment. This is evident in both the smallest freshwater swamps, found in areas such as Nee Soon catchment in Singapore (Ng and Lim, 1992) to extensive floodplains, like the 15,000 km2 floodplain of Tonle Sap (Cambodia) that is inundated during the flood pulse of the Mekong river (MRC, 2005, cited in Arias et al., 2012). River processes drive heterogeneity within the floodplains. For example, the mosaic of rainforest floodplain communities of the Fly River are associated with specific geomorphic features on the floodplain, such as scroll complexes, backswamps, alluvial plains of minor tributaries, blocked valley swamp and lakes (Blake and Oliver, 1971, cited in Dietrich et al., 1999). Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 722 Progress in Physical Geography 38(6) Figure 3. Extensive mid-channel island on the anabranching lower Irrawaddy, downstream of Pyay, Burma (B3). Periodically inundated woody vegetation grows at the highest elevations (left) and the scroll bar comprises seasonally inundated vegetation and a small backswamp (centre). While forested floodplains dominate within the region, natural, non-forested floodplains do occur in the lower basins of some of the major rivers, such as the Irrawaddy, Chao Phraya and Mekong (Corlett, 2009), as exemplified in Figure 3. The biota of both the river and the associated floodplains and wetlands shows an ecological association with the flood pulse, although this is very under-researched (Dudgeon, 2000a). For example, seasonal patterns of phytoplankton and zooplankton are related to the monsoonal flood pulse, with a high abundance in the dry season and a low abundance in the wet season (due to washout, dilution and high turbidity), although interstream variation in response has been observed (Dudgeon, 2000a). Fish abundance is also associated with the flood pulse, with generally higher numbers during stronger monsoons. Along the Mekong River, life cycles are synchronised with the flow regime, with upstream breeding migrations occurring during the wet season and downstream migrations occurring when water levels recede (Welcomme, 1979). Similarly, the inundated floodplains in Bangladesh provide important nurseries for larvae and juvenile fish species and fish abundances are positively correlated with flood extent (De Graaf, 2003). Indeed, in areas with two monsoonal pulses (e.g. south India), fish may reproduce twice (Dudgeon, 1992). Ecological associations with the flood pulse are not restricted to aquatic organisms either; many species of deer that occur in Asia (e.g. Hydropotesinermis, Elaphurusdavidianus, Axis porcinus) use or are even confined to the seasonally inundated floodplains and riparian wetlands and some show adaptations such as splayed or very large hooves for grazing these habitats (Dudgeon, 2000a). The increased discharge associated with the flood pulse also influences water chemistry, such as ion concentrations, salinity, turbidity and dissolved organic matter, which is likely to have a significant ecological impact, although this remains under-researched (Dudgeon, 1992). While some associations between hydrology and ecology are described, links between fluvial processes and the vegetation of these systems have received very little research attention, despite the fundamental functional importance of vegetation within river systems that has been acknowledged elsewhere (e.g. Corenblit et al., 2009a, 2009b). Existing studies have tended to be piecemeal and site-specific, but these are synthesized here from two perspectives: the role of hydrological and geomorphological processes on vegetation and the active role of vegetation in modifying physical processes. III Interactions between hydrology, geomorphology and vegetation in Southeast Asian rivers and floodplains 1 Hydrological and geomorphological impacts on riparian vegetation Literature on riparian and wetland vegetation within the region has been heavily focused towards botanical descriptions without strong associations with physical processes. However, these descriptions do show linkages between Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 Moggridge and Higgitt 723 vegetation and the flood pulse. Research on the floodplains of the very large rivers is sparse. Many of the largest rivers exhibit anabranching planforms, which are some of the most complex and least understood systems (Latrubesse, 2008), and while research on floodplain vegetation of the large, anabranching systems in the Neotropics has shown complex mosaics of floodplain wetlands associated with hydrological and geomorphological processes (e.g. Latrubesse, 2012; Stevaux et al., 2013; and previously cited examples) only a few isolated studies exist within the study region. A study by Arias et al. (2012) on Tonle Sap Lake (part of the Mekong system) drew linkages between the mosaic of floodplain vegetation types, which included gallery forest (i.e. riparian forest that occurs in an environment that is otherwise unable to support forest), flooded shrubland and open water, with the duration of inundation in wet, normal and dry years. Similarly, although peripheral to the study region, vegetation within the active zone of the braided Brahmaputra (category B2 in Table 1) shows a succession with elevation (and thus inundation): short seasonal grass was found 0.5–2 m above bar height, taller grass and shrubs were found at 2.5 m, and 4 m high grass and trees were found up to 8 m above bar height (Sarma, 2005). Observation of vegetation on an in-channel island on the anabranching Irrawaddy (B3) also showed a distinction between vegetation type according to inundation regime (Figure 3). More detailed research has been conducted on smaller meandering systems within the region. The geomorphic complexity of the floodplain of the meandering sections of the Fly River (B1) (Papua New Guinea) shows specific vegetation assemblages with different landforms and is comprehensively described by Rau and Reagan (2009). Open canopy forests, composed of Pometia, Terminalia, Alstonia, Planchonia, Bischofia, Cananga and Nauclea are found on scroll bars and crevasse splays, while Campnosperma, Terminalia, Syzygium, Nauclea and Myristica dominate in the lower-elevation backswamps of the system. In parts of the floodplain that are permanently inundated, open swamp woodland dominated by Melaleucacajuputi or Naucleaorientale or aquatic grassland species occurs (see Rau and Reagan, 2009, for full species list). Similarly, vegetation communities on the monsoonal Adelaide River floodplain (B1) in Australia showed distinctions along a seasonally flooded elevation gradient: Syzgium spp. and Nauclea spp. dominated in areas not flooded by more than 1 m, while Melaleuca spp. dominated in areas flooded by 2 m of water (Bowman and McDonough, 1991). These studies exemplify the rich diversity of vegetation that can occur within floodplain environments, but this is generally unrecorded in the region. Other botanical descriptions in the region have mainly been confined to small swamp forests, without information on the river system. For example, Yamada (1997) describes the vegetation composition of freshwater swamp in Burma and showed that areas that differ in inundation have distinct tree communities; patches that were only inundated during the monsoon were dominated by Albiziaprocera and Buteafrondosa, among others, while continuously inundated areas that only dry out for a few weeks per year were dominated by Xanthophyllum spp. and Dalbergiareniformis. Similarly, in the small, equatorial catchment of Nee Soon in Singapore, forest composition was related to the magnitude of water-level fluctuation and the intensity of disturbance (Corner, 1978). Within the freshwater swamp of Nee Soon, forest community composition varied with flood extent: Palaquiumxanthochymum dominated in areas that were frequently flooded; Xylopiafusca dominated at higher elevations that were less prone to inundation; and Lophopetalummultinervia, Cratoxylumarborescens and Alstoniaspathulata occurred adjacent to river channels, where physical disturbance was greatest. A similar relationship between elevation and vegetation composition is Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 724 Progress in Physical Geography 38(6) described in Kalimantan by MacKinnon et al. (1996). Many of the trees observed in these environments have adaptations, such as stilt or buttress roots or pneumatophores, to tolerate inundation (Page et al., 1999). Corlett (2009) suggests that high flows in riparian zones may facilitate plant dispersal (and thus play a significant role in the life cycles of those species), but also adds that studies on this are absent. In Borneo, the dominant tree of freshwater swamp, Dryobalanopsrappa, can reproduce vegetatively following mechanical disturbance, which may play an ecological role in maintaining the tree populations within the swamp, by complementing seed regeneration (Yamada and Suzuki, 2004). While this process requires significant further research, the observation of a complementary role of sexual and asexual reproduction in floodplain trees has synergies with the Salicaceae in northern temperate floodplains. Asexual reproduction is important in maintaining Salicaceae populations, as seed establishment is tightly coupled to flow regimes and thus exhibits high interannual variability (Barsoum, 2002; Mahoney and Rood, 1998). While there has been no research into the life cycle of floodplain vegetation in Southeast Asia, observation has shown that significant variation from ‘average’ hydrological conditions can have an adverse effect. For example, vegetation on the floodplain of the Mekong is adapted to a ‘normal’ wet season and substantially longer floods can lead to die-off and rotting, which can contribute to de-oxygenated conditions in the system (Welcomme and Halls, 2004). The above case studies all demonstrate the importance of lateral connectivity between rivers and floodplain systems for vegetation communities. Studies on other taxa (discussed in the previous section) further confirm the ecological importance of these exchanges. Following a synthesis of tropical Asian river ecology focusing on fish and invertebrates, Dudgeon (1992) asserts the ecological significance of these lateral exchanges, suggesting that longitudinal models such as the River Continuum Concept (RCC) (Vannote et al., 1980) present an incomplete description of these processes. Further to this, research from the Neotropics has suggested that the Flood Pulse Concept may be too simplistic for very large floodplain systems: water may enter the floodplain from a variety of pathways, including rainfall, flooding of local tributaries, groundwater and main channel exchanges, which interact with the morphological complexity of the floodplain to create a mosaic of different hydrological conditions (Mertes et al., 1995). This is concurrent with conceptual advances from temperate regions, where frameworks have extended from a continuum approach and consider river systems as mosaics of heterogeneous, dynamic patches, encompassing lateral, longitudinal and vertical exchanges (e.g. Frissell et al., 1986; Poole, 2002) and geomorphological processes (Thorp et al., 2006). Such models may be useful frameworks for Southeast Asian systems, but this has not been tested to date. 2 The influence of vegetation on hydrological and geomorphological processes The extensive variation in the size of rivers and floodplains within Southeast Asia makes any conceptual generalization of the active role of vegetation problematic. Much existing regional work in this field has been at the catchment scale, where research has concentrated on the impacts of large-scale deforestation, where increases in river flow, stormflow, sediment load and decreases in water quality and nutrients have been observed following vegetation removal (see Douglas, 1999, for a comprehensive review). In very large systems, vegetation is unlikely to have a major direct geomorphic role within the active zone, but large changes in vegetation within the catchment can still be hydrologically and geomorphologically significant. For example, within the Neotropics, studies of the Araguaia river catchment in Brazil Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 Moggridge and Higgitt 725 Figure 4. Fallen trees, perpendicular to flow direction, create small jams and form step pools in the Sungei Melinau Paku, Gunung Mulu National Park, Sarawak, Malaysian Borneo. (catchment area 375,000 km2, mean annual Q ¼ 6500 m3 s-1), where 25% of the catchment has been deforested, showed a 25% increase in mean annual discharge from pre-deforestation levels (Coe et al., 2011) and a 31% increase in bed load transport (Latrubesse et al., 2009). This has initiated a significant morphological response: gullies have formed in the upper reaches of the river, the number of islands in the channel has decreased by 30%, the number of sandy bars has risen substantially and the channel planform has shifted from anabranching to braided (Latrubesse et al., 2009). Thus, vegetation still exerts a strong active control in very large systems. Within temperate systems, riparian vegetation has been shown to play an active role in altering flow and sediment dynamics and influencing river planform (e.g. Gurnell et al., 2012). Studies within Southeast Asia describe vegetation growing on braid bars (e.g. Brahmaputra, Sarma, 2005; Fly, Pickup and Marshall, 2009) and anabranch islands (e.g. Mekong, Gupta and Liew, 2007, and observation of the Irrawaddy, shown in Figure 3), but the potential active role of this vegetation has not been explored. A collection of studies from the region do demonstrate an active role of vegetation, but this is confined to the channel scale and is thus only relevant to small river systems. At a patch level, work has shown that riparian vegetation alters the light, temperature and hydraulic conditions within small river channels, which influences the invertebrate communities present (Dudgeon, 1989). Similarly, a study on the habitat requirements of fish on the floodplain wetlands of the meandering Mulgrave River in northern Australia (B1) (catchment area 810 km2, mean annual Q ¼ approx. 50 m3 s-1) showed that the riparian and in-channel vegetation supported the fish communities by creating habitats which changed seasonally with the flood pulse to meet the different requirements of the fish at key stages of their life cycle (Rayner et al., 2008). Large wood deposited within river channels can be an important influence on physical processes. For example, Figure 4 shows fallen trees across a small channel in Sarawak, Borneo, which have created a small jam and altered flow patterns. Published work within the region also demonstrates the importance of large wood. A study of three small headwater streams in Sabah, Malaysia (catchment area 0.5–10 km2), showed that in small channels (i.e. a channel width around 20 m where the channel can be blocked by a single fallen log), tree fall was a major influence on channel morphology and processes (Spencer et al., 1990). Trees changed the longitudinal profile of the channels by creating organic steps, with falls of 2–3 m in height. The rate of rotting of the logs, lateral movement of the channel and the magnitude of high flows influenced the residence time of these jams, which the authors suggest may be lower than for temperate systems. Nonetheless, these jams may still be morphologically important; in addition to the direct changes to the channel, the jams collect and store sediment in minor and moderate events and release it in major and extreme events, increasing the importance of high-magnitude, low-frequency events for the Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 726 Progress in Physical Geography 38(6) removal of fluvial sediment. These findings are supported by observations of headwaters in the forested Bukit Tarek catchment in Penninsula Malaysia (A1) (catchment area 0.33 km2), where 500–600 large wood accumulations per ha of channel were observed, storing 20–40 m3 ha sediment (Gomi et al., 2006). Within the typically nutrient-poor river systems of Southeast Asia, large wood can also play an important role in retaining organic matter. Study of Neotropical streams has shown that reduced hydraulic forces behind dams and the accumulations of organic detritus provides important shelter and food resources for a range of animals, which promotes biodiversity (Wantzen and Junk, 2000). In a global synthesis of the role of wood in river systems, Gurnell (2013) compared the storage of wood in tropical rivers, derived from a small collection of studies from Central and South America, to forest types in other ecoregions. Wood storage in rivers in tropical rainforest (mean 189 m3 ha-1, median 144 m3 ha-1) was significantly less than for redwood or conifer forest rivers, but significantly greater than deciduous and Salicaceae dominated rivers. In this comparison, Gurnell (2013) identifies two factors which drive contrasts between wood processes in different systems: (1) the wood budget, i.e. the relative contribution and key processes governing wood input, retention and output within systems; (2) the ability of the wood to regenerate. While the conclusions of Spencer et al. (1990) from Borneo have not been tested further within Southeast Asia, studies from the wider tropics do suggest that further research within this region could make an important contribution to this field. For example, in an overview of tropical rainforests, Walsh and Blake (2009) suggest that rainforest vegetation may exert a greater role on channel processes than in temperate zones and identify this as a defining characteristic of tropical rivers. Further, Cadol and Wohl (2010) and Cadol et al. (2009) make comparisons of wood budgets between tropical and temperate river systems, based on field studies in Costa Rican small headwater streams (catchment area 0.1–8.5 km2). The studies showed that wood in the channel was transported from upstream sources rather than occurring from local recruitment (i.e. tree fall into the channel), which differed to local recruitment-dominant temperate systems. Reduced wood loads and lower retention times (mean residence time for a piece of wood was 4.9 years) in tropical rivers compared to temperate systems were also observed, which the authors attribute to flashier flow regimes, the branched structure of the trees and higher decay rates in tropical environments. They also infer that this may reduce the long-term geomorphological importance of wood jams in these systems, which opposes Walsh and Blake’s (2009) assertions and highlights the need for further research in this field. Synergies may be drawn between Southeast Asian and Costa Rican systems, although the relative importance of recruitment processes may differ in Southeast Asia from what Cadol and Wohl (2010) and Cadol et al. (2009) observed, on account of the variation in storm and hurricane frequency and intensity across the region. Further to observations on the wood budget in the Neotropics, Wantzen and Junk (2000) found that large wood had the ability to regenerate roots and twigs, which further alters flow patterns within the channel. While these observations are not within the study region, it is entirely plausible that trees within Southeast Asia may exhibit the same characteristics. There is an acknowledged bias in large wood studies from the temperate zone and a need to expand understanding into different regions and different river types (Collins et al., 2012; Gurnell, 2013). Existing studies from tropical rivers do suggest a different morphological behaviour of wood, but further work on Southeast Asian river types, with different flow regimes and sediment dynamics, could extend understanding further. Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 Moggridge and Higgitt 727 The studies discussed above demonstrate the geomorphological importance of in-channel vegetation. A study by Iwata et al. (2003) on the headwaters of Rayu River in Malaysian Borneo (discharge of study reaches ¼ 0.01–1.17 m3 s-1) extends this by demonstrating that living riverbank vegetation can influence in-stream habitat. The research investigated the impact of deforestation on invertebrate communities, but included field observations of stream habitat next to primary and secondary riparian forest. Channels bordered by primary riparian forest were characterized by coarser substrate, stable banks, a greater proportion of erosional habitat and less channel cover, while channels with secondary riparian forest had finer substrate, more eroded banks, more depositional habitat and more tree cover on the water. This suggests a reach-scale influence of particular vegetation communities on sediment dynamics, but the landform-scale morphological importance of this is not considered (as it was not an objective of the study) and more detailed, directed studies are needed to confirm and extend these findings. This synthesis of existing studies demonstrates that vegetation has an active role in influencing geomorphological processes at the channel and reach scale within the smaller river systems in the region (such as type A1 identified in Table 1). Work on in-channel large wood in the wider tropical region indicates that processes of wood entrainment, storage, geomorphic influence and decay differ to temperate systems, but further work in Southeast Asia is required. Research to date has mainly focused on vegetation within the river channel; the active role of vegetation on inchannel landforms, such as bars and islands, and on floodplains and their wetlands has not been explored, despite studies in temperate systems that show the critical geomorphological importance of this (e.g. Corenblit et al., 2009b; Gurnell and Petts, 2006). This raises a number of pertinent research questions for the region, which are discussed in the following section. IV Directions for future research There is a clear deficit in understanding of the interaction between vegetation and fluvial processes in Southeast Asian rivers and wetlands, which is reflective of a wider absence of fluvial geomorphological research within tropical systems (Walsh and Blake, 2009). In 1990, the urgent need for the most basic descriptive and inventory information on forested wetlands within the region was recognized by Jusoff and Majid. Subsequent progress has been very slow (although see, for example, Yamada, 1997), which highlights the extent of the knowledge gap. Existing descriptions of floodplain vegetation have shown a relationship between species distributions and variations in the flood pulse, which demonstrates the ecological importance of inundation and connectivity in tropical floodplain systems and is synonymous with existing concepts (e.g. Flood Pulse Concept, Junk et al., 1989) and field studies from temperate environments (e.g. Auble et al., 1994; Hupp and Osterkamp, 1985; Johnson, 2000). However, these studies lack a process-based explanation for the observed vegetation distributions, which raises important questions for future research. Although the influence of water levels is acknowledged, the ecological importance of particular characteristics of the pulse (as researched in Neotropical rivers and described by Welcomme and Halls, 2004) is not understood. Further, with the exception of Rau and Reagan’s (2009) work on the Fly River in Papua New Guinea, studies from Southeast Asia have not been extended to consider hydrogeomorphological processes. Current research from temperate systems (e.g. Stanford et al., 2005; Steiger et al., 2005) and the Neotropics (e.g. Marchetti et al., 2013) has progressed from the Flood Pulse Concept, suggesting that simple linear gradients of inundation are insufficient in explaining vegetation patterns; vegetation patterns reflect hydrogeomorphological processes Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 728 Progress in Physical Geography 38(6) which create heterogeneous and dynamic habitat mosaics within the river landscape. The applicability of these concepts to Southeast Asian systems is an important and worthwhile area for future research. Boulton et al. (2008) rightly caution against an uncritical extrapolation of temperate models into tropical rivers and any comparative work must be supported by robust field evidence. This is particularly important when the large anabranching systems are considered; the majority of these occur within the tropical latitudes (Latrubesse, 2008) and with most models of vegetation dynamics developed through research on smaller meandering or braided systems within temperate zones (e.g. Stanford et al., 2005; Steiger et al., 2005) such models may be inappropriate to these systems. Similarly, while some of the findings on vegetation in the large meandering and anabranching systems of the Neotropics may be relevant to equivalent systems within Southeast Asia, the different flow regimes and species within the Southeast Asian systems may also create important differences which could extend understanding in this field. Thus, there is a need to explore these processes across all different types of river identified in Table 1 (and indeed in river types that might not have been considered), in systems of different size, flow regime and with different vegetation communities. Understanding the ecological requirements of riparian and wetland species and the association with physical processes at each stage of their life cycle is also fundamental. These processes are well researched in temperate floodplains where models of vegetation, flow and sediment interactions are applied to floodplain management and conservation (e.g. Mahoney and Rood, 1998; Richards et al., 2002). However, they are not currently understood in the exceptionally biodiverse Southeast Asian systems, where such models could effectively support conservation in the region. Research is needed on reproduction and initial establishment of the dominant floodplain species and the significance of the flood pulse and associated disturbance on influencing these processes. There are indications from the literature that vegetation actively influences fluvial processes in Southeast Asian systems, as evidenced predominantly from studies of large wood within small river channels. This specific field has been well researched in temperate systems and there is a widely acknowledged need for studies in other ecoregions (Collins et al., 2012; Gurnell, 2013; Montgomery and Piégay, 2003). As outlined in the previous section, there are notable contrasts in wood processes between river systems and further research in Southeast Asia could make an important contribution to global knowledge in this field, due to the different species, disturbances and wood decay rates (Spencer et al., 1990). Research on the role of wood in the medium and large systems is also needed, as this has not been considered at all in tropical systems. The active role of vegetation in influencing fluvial processes extends beyond the morphological changes within the channel (Gurnell et al., 2012) and broad catchment-scale processes. Research outside of the tropics has shown vegetation to have a major influence on fluvial morphology and processes across the floodplain, but no existing research on this within Southeast Asian systems or the wider tropics was found in this synthesis. There is a pertinent need to research how vegetation influences landform growth, development and residence time at different scales and across river types in the region (e.g. B1B3). Many of the very large rivers within the region have established vegetation within the active zone and the physical role of this needs researching, as there have been very few studies on large systems. Across all of the river types and catchment sizes within the region, the different species found in tropical floodplains and wetlands, with their faster growth and decay rates, coupled with the flow regimes and high sediment loads of these systems, could provide new, fresh insights and Downloaded from ppg.sagepub.com at PENNSYLVANIA STATE UNIV on May 13, 2016 Moggridge and Higgitt 729 potentially extend existing temperate-focused models such as Corenblit et al. (2009a). With tropical catchments covering the largest area of the Earth’s climatic zones, there is a need for more representative research on these processes, rather than broad extrapolation of temperate models into the area (Boulton et al., 2008). Southeast Asian systems are inherently very variable, both in river types and in the large variation in size. Thus, it is problematic to make wide generalizations about the interaction of vegetation and fluvial processes and to what extent existing models can be appropriately applied. Boulton et al. (2008) also recognize this and suggest that the extrapolation of (ecological) models may not be an issue of temperate against tropical systems, but that the model needs to be applied to an appropriate and comparable system. Similarly, direct comparison between tropical and temperate systems does not necessarily have to be the focus for further research; empirical data on vegetation and fluvial processes must be the priority and this may even yield new models in addition to advancing global knowledge of these processes. This supports the argument made by Douglas (1993: 104) for a greater focus on research within tropical regions across the discipline of geography: ‘it is high time the whole world sees the tropics as the ‘‘norm’’’. Vegetation and hydrogeomorphological processes provide the physical habitat that underpins all the other biotas in this exceptionally biodiverse environment. It supports the functional integrity of the ecosystem and thus its long-term sustainability. The environment within this region is undergoing rapid rates of loss and human alteration (Dudgeon, 2000b), with few rivers of Southeast Asia still in a natural state (Gupta, 2005). Therefore, in addition to the academic value of this research, there is an urgent need to understand these systems, to support and inform effective conservation and management. Acknowledgements H. Moggridge would like to thank the University of Sheffield for funding this fellowship and the National University of Singapore for their kind hospitality. Thanks also go to Paul Coles (University of Sheffield) for his assistance with the production of Figure 1 and to the two anonymous reviewers for their helpful and constructive comments. 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