The Fourth Dimension: an Overview of Altitudinal Migration

The Fourth Dimension: An Overview of Altitudinal Migration

Presented by M.Sc. Stefan Kreft Fundación Amigos de la Naturaleza, Santa Cruz, Bolivia “Migratory Workshop Species: Linking 25th anniversary Ecosystems of the Bonn and Convention Disciplines“

© Stefan Kreft rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 e community and population level ing altitudinal movements and longi-latitudinal migration + Proximate factors influenc + Important differences between altitudinal migration + Important conservation implications + Altitudinal migration and CMS S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal Overview topics: + The fourth dimension: Phenomena, perceptions and definitions 25th Annive Animal groups,+ regions and ecosystems involved + Spatio-temporal dimensionality + movements on th Altitudinal ...... escape movement local movement

rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 hilltopping intratropical migration movement disper sal

migration displacement nomadism x No standard (widely accepted) definition exists(?)! Variety of terms used: Phenomena, perceptions and definitions – setting the stage...

S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive elevational altitudinal vertical – a In studies, ad-hoc few definitions are given, in others definitions are implicit, in the majority they appear not to have been contemplated consciously... rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 volant animals: birds etc. organisms, benthic mesopsammon Interesting, but too heterogeneous!

S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive -> Daily vertical migration of plankton in oceans and lakes migration of soil -> Vertical -> Change in flight altitude of The act of moving from one spacial unit to another, but always including a movement in vertical dimension. the This would include, for instance: In a DEFINITION complementary to Baker’s (1978) very broad concept of migration, “altitudinal migration” might be taken as: .

rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 mountainous area including a change in altitude in a regular to-and-fro movements, always A possible, still broad definition: Animal populations or parts of these show

S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive Altitudinal migration in taxa other than birds:

Insecta: Lepidoptera Costa Rica Haber 1993, Stevenson & Haber 2000*, Haber & Stevenson in prep. California Shapiro 1986 Kenya Awodey 1978 Hymenoptera: Costa Rica Hunt et al. 1999* Vespidae Hymenoptera: Japan Tomono & Sota 1997 Apidae Diptera Japan Arakawa et al. 1991*, Kurahashi et al. 1991* Coleoptera China Xu & Liu 1998 ... Pisces Salmonidae Europe, N-America ... Mammalia Chiroptera Costa Rica Timm & LaVal 2000* Galápagos Islands (Ecuador) McCracken et al. 1997 Mexico Montalvo 1997 Chile Sanborn & Crespo 1957 Primates Sumatra (Indonesia) Buij et al. 2002 China Fooden 1986 Perissodactyla Costa Rica Lawton 2000* Artiodactyla Rocky Mountains (USA) Gruell 1958, Moisan 1958, Shaw 1958, Robel 1960, Hebert 1973, Oosenbrug & Theberge 1980, Morgantini & Hudson 1983 Himalaya (India) Sarkar et al. 1999 Japan Takatsuki et al. 2000 Southern Alps (New Zealand) Tustin & Parkes 1988 Alps (France) Géroudet 1972*, Rochat 1996 Alps (Italy) Parrini et al. 2003 Pyrennees (Spain) Herrero et al. 1996 Norway Albon & Langvatn 1992, Mysterud 1999 Carnivora: Spain Caussimont et al. 1993 Ursidae Carnivora: Rocky Mountains (USA) Logan & Irwin 1984 Felidae

* : entire publication revised (others just through title and keywords) Ecosystem used by altitudinal migrant birds:

Ecosystem# Studies (n=116)* Diverse 15 Temperate alpine open land/subalpine shrubs 7 Temperate montane conifer forests 8 Temperate montane mixed forests 6 Temperate montane broadleaf forests 5 Puna/Páramo 12 Tropical arid/semi-arid lowlands/tablelands 11 Polylepis woodland 1 Tropical humid forests 67 Subtropical humid/semi-humid montane forest 4 * multiple counts possible Spatio-temporal dimensionality of altitudinal migration

resident partial partial partial partial partial total migrant migrant migrant migrant migrant migrant upper limit of breeding area Altitude

lower limit of breeding area

area occupied by population throughout annual cycle area occupied only outside breeding season area occupied only during breeding season Studies of altitudinal migration of birds in South America

Single or few spp., raw data

Few spp. in detail or number of spp., but raw data Number of spp. in detail

Entire guild or family in detail

Community-based, detailed

Sonntag, 20. Juni 2004 (4).jpg after breeding 2: return migration (upslope) rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 Non-breeding season 1: outward (downslope) migration

After breeding - Nov

Breeding Sept of Neotropical birds: ⌫

2 July Return migration upslope Return migration Outward migration in the majority downslope

May

Direction and timing of altitudinal migration : E.g., on eastern (Amazonian) slope of eastern Andes in Bolivia: Mar Non-

1

⌫ breeding

S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive Jan Precipitation Monomodal precipitation cycle: Direction Timing: - daily in high numbers! 9

rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 Schaefer 1953 Benson 1982 Rosselli 1994 Cody 1970 Rabenold & Rabenold 1985 Saniga 1995 Stiles 1985 Stiles 1985 Raman 19 Cody 1970 Cody 1970 Cody 1970 Powell & Bjork 1995 Robinson etal. 1988 Martín-Vivaldiet al. 1995 Dixon& Gilbert 1964 Laymon 1989 Stiles 1985 Stiles 1985 Blake& Loiselle 2002 Loiselle& Blake 1991 Horvath &Sullivan 1988 Monroe 1968 Géroudet 1972 Oatley 1966 Voisin 1995Pearson 1974 Pearson 1974 Hughes 1970 Hughes 1970 Hughes - 1970 daily! altitudinal interval [x 100m]-> [x interval 2altitudinal 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 Venezuela, Coastal Range Range Coastal Venezuela, Tanzania, Uzambara Mts. Uzambara Tanzania, slope Carribbean Costa Rica, Chile, Pacific slope Andes Pacific Chile, North Carolina, Appalachians Carolina, North Slovakia, Carpathians Carpathians Slovakia, Costa Rica, Carribean slope Carribean Costa Rica, Costa Rica, Carribean slope Carribean Costa Rica, Western Ghats India, Chile, Pacific slope Andes Pacific Chile, Chile, Pacific slope Andes Pacific Chile, slope Andes Pacific Chile, central Costa Rica Costa Rica central Peru, Andes Spain, SierraNevada Mts. Rocky Utah, California, Sierra Nevada slope Carribean Costa Rica, Honduras Costa Rica, Carribean slope Carribean Costa Rica, slope Carribean Costa Rica, slope Carribean Costa Rica, Mts. Rocky Arizona, Alps Italy, France, Alps France, slopeAndes Pacific Peru, slopeAndes Pacific Peru, slopeAndes Pacific Peru, slopeAndes Pacific Peru, South Africa slopeAndes Pacific Peru, ) catchers y ccentors) ccentors) (Pycnonotidae) rannidae, New World fl World New rannidae, (Falconidae, falcons) (Falconidae, y (Phoenicopteridae, flamingos) (Phoenicopteridae, (Trogonidae, quetzals and trogons) trogons) and quetzals (Trogonidae, (T (Trochilidae, hummingbirds) hummingbirds) (Trochilidae, (Corvidae, crows) crows) (Corvidae, (Pipridae, manakins) manakins) (Pipridae, (Tichodromidae, wallcreepers) wallcreepers) (Tichodromidae, (Turdidae, thrushes) thrushes) (Turdidae, (Nectariidae, sunbirds) (Nectariidae, (Trochilidae, hummingbirds) (Trochilidae, (Carduelidae, cardueline finches) cardueline (Carduelidae, (Threskiornithidae, ibises) (Threskiornithidae, (Trochilidae, hummingbirds) (Trochilidae, (Trochilidae, hummingbirds) hummingbirds) (Trochilidae, (Strigidae, owls) owls) (Strigidae, (Turdidae, thrushes) thrushes) (Turdidae, a (Prunellidae, (Emberizidae, New World warblers) World New (Emberizidae, (Tyrannidae, New World flycatchers) World New (Tyrannidae, (Emberizidae, New World sparrows) sparrows) World New (Emberizidae, (Cathartidae, New World vultures) World New (Cathartidae, (Laridae, gulls) (Laridae, (Thraupidae, tanagers) (Thraupidae, (Paridae, chickadees) (Paridae, (Tyrannidae, New World flycatchers) World New (Tyrannidae, (Pipridae, manakins) (Pipridae, (Tyrannidae, New World flycatchers) World New (Tyrannidae,

S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive Tersina viridis Nectarinia olivacea Nectarinia Corapipo leucorrhoa Corapipo frontalis Muscisaxicola M. flavinucha M. Junco hyemalis Junco Tichodroma muraria Elvira cupreiceps Lampornis hemileucus madagascariensis Hypsipetes Species (Family) Country, mountain range Source M. albilora M. maculirostris M. mocinno Pharomachrus Turdus nigriceps Turdus collaris Prunella Parus gambeli Strix occidentalis Discosura conversii Discosura delphinae Colibri pipra Pipra spp. various phaenotus Junco psaltria Carduelis spp. various natalensis Cossypha Pyrrhocorax graculus Pyrrhocorax Plegadis ridgwayi Larus serranus chilensis Phoenicopterus gryphus Vultur megalopterus Phacoboenus Altitudinal intervals crossed rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004

distance traveled [km]-> traveled distance Powell & Bjork 1995 Source Horvath & Sullivan 1988 Stiles& Clark 1989 Loiselle& Blake 1991 Laymon 1989 Benson 1982 Benson 1982 Benson 1982 Benson 1982 Dowsett (1988)

L a y m o n

1 9 8 9 central Costa Rica Costa Rica central Arizona, Rocky Mts. Rocky Arizona, slope Carribean CostaRica, slope Carribean CostaRica, California, SierraNevada southern Africa southern Africa southern Africa southern Africa Zambia Malawi, - (Trochilidae, hummingbirds) (Trochilidae, (Motacillidae, pipits) pipits) (Motacillidae, (Trogonidae, quetzals and trogons) trogons) and quetzals (Trogonidae, (Turdidae, thrushes) (Turdidae, (Turdidae, thrushes) thrushes) (Turdidae, sunbirds) (Nectariniidae, (Strigidae, owls) owls) (Strigidae, (Emberizidae, New World warblers) World New (Emberizidae, (Sylviidae, Old World warblers) warblers) World Old (Sylviidae, Spotted owl S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive Strix occidentalis Distances traveled: Junco phaenotus Junco Species (Family)Pharomachrus mocinno Country, mountain range Microchera albocoronata Microchera spp. various Strix occidentalis torquata Saxicola scita Stenostira novaeseelandiae Anthus cuprea Nectarinia stellata Pogonocichla Altitudinal migration on the community level:

Chesser (1997) Bolivian Andes 15 probable altitudinal migrants Tyrannidae (New World flycatchers) 26% n=57 20% Stiles (1988) 69 seasonal migrants Caribbean slope 19 daily movers 6% “Resident” forest birds 17 movers of uncertain status n=345 5%

Hilty (1997) Colombia, Pacific slope western Andes 57 altitudinal migrants Cloud-forest birds 21% n=271

Thiollay (1980) Nepal, Himalaya 173 altitudinal movers Montane breeding birds 51%! n=340 Trophic guilds:

High coincidence in studies about feeding guilds amongst altitudinal migrants of tropical humid forest (by order of importance):

1. Nectarivores Daily movers: 2. Frugivores Strong-flying birds of prey, aerial insectivores

Habitat preference:

Most Neotropical altitudinal migrant birds are forest-based. rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 distributional core area or leave to season, young birds are frequently (e.g., Loiselle & Blake 1991). Diamond 1973, own observations). but partial migration seems to be more common underinvestigated (Ketterson & Nolan 1976). respect to migration altitudinal (e.g., Chesser 1997). • of Cases differential migration are known, altitudinal but

S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal Subspecies or populations may differ inbehaviour with their 25th Annive Altitudinal migration on the population level: • • Particularly after the breeding find their own territory (dispersion). They stay in the periphery (e.g., • as migratory partially There are well migratory populations, as totally expulsed from the most favourable rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 (arthropods, fruits, vertebrate prey ricanes, natural fires) - semi-predictable - semi-stochastic -~predictable

S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive Proximate factors influencing altitudinal movements: etc.) - ~predictable -> catastrophic events (hur -> adverse weather events (heavy rains, snow storms) -> Phenologies of food resources -> Competition (intra--> interspecific) and . . . . . interspecific competition areas ~linear

rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 areas disjunct . areas ~visually connected conical shape of mountains at breeding and non-breeding areas. . broad-front migration weather events ~the same S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive ...limits of breeding area often defined ~by ... Other than in longi-latitudinal migration...... breeding and non-breeding distribution ~restricted ... extreme ... in... many cases especially breeding Important differences between altitudinal migration and longi-latitudinal migration: Geometry rules...... especially breeding (but also non-breeding) ...breeding areas ~restricted by ~ rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 intain populations of stributional limits New calculations of actual distributional areas required Complementary, year-round protection necessary, in both breeding and non-breeding area Migration corridors Hunting, capture for pet market (e.g., cage bird traders in Brazil!) southeast Nectarivores and frugivores important mobile links (Gilbert 1980): ma plants and their di Climate change: (altitudinal) migrants depend on multiple areas with independent phenological rhythms; rapid change may lead to a miscoordination between the breeding- non- and breeding areas ! ! ! ! ! ! S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive Some important implications for conservation: ”Cycle” criterion Varying percentage of altitudinal migrant individuals within a species >

rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 ”Border-crossing” criterion Semi-stochastic/-predictable Semi-stochastic/-predictable ? ) some altitudinal migrants travel > events not included > ( as far asas far 100 km, but the majority only short distances X Altitudinal migration and the CMS: ”100 km” criterion But consider, e.g., Nepal/India, France, Zimbabwe/Moçambique ”Predictability” criterion x? ”Significant proportion” criterion S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive But include daily movers and escapers! Guatemala/Belize, Switzerland/ Italy, > (applied only by GROMS): rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 behaviour. CMS may look for synergisms with altitudinal conservation corridors! and ecologically important migratory altitudinal as well “CMS migrants“! local/regional activities for conservation of => Convincing argument for the need S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive Altitudinal migration represents a widespread Powell & Bjork (2004) )

rsary Bonn Convention -rsary Bonn Convention Berlin, 22./23.06.2004 Procnias tricarunculata ( the case of Three-wattled Bellbird

S. Kreft: Altitudinal Migration Migration S. Kreft: Altitudinal 25th Annive Altitudinal and long-distance migration linked: Acknowledgments: PD Dr. Hermann Ellenberg - Federal Research Institute for Timber and Forestry, Hamburg, Germany Dr. Ralf Strewe – Fundación Pro-Sierra Nevada de Santa Marta, Santa Marta, Colombia Lic. Natalia Araujo - Fundación Amigos de la Naturaleza, Santa Cruz, Bolivia Dr. Klaus Riede – GROMS at Alexander Koenig Research Institute and Museum of Zoology, Bonn, Germany

& too many more to be mentioned here...

Research funded by: Hamburg Grant for Graduate Students German Academic Exchange Service German Society for Tropical Ornithology & my parents, Rosemarie & Horst Kreft