Kleptoparasitism of Shoebills Balaeniceps Rex by African Fish Eagles Haliaeetus Vocifer in Western Tanzania
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Tanzania Journal of Science 45(2): 131-143, 2019 ISSN 0856-1761, e-ISSN 2507-7961 © College of Natural and Applied Sciences, University of Dar es Salaam, 2019 Kleptoparasitism of Shoebills Balaeniceps rex by African Fish Eagles Haliaeetus vocifer in Western Tanzania Jasson RM John1* and Woo S Lee2 1Department of Zoology and Wildlife Conservation, College of Natural and Applied Sciences, University of Dar es Salaam, P. O. Box 35064, Dar es Salaam, Tanzania. 2Department of Forest Sciences, College of Agriculture and Life Sciences, Seoul National University, Seoul 08826, Korea. *Corresponding author E-mail: [email protected] Abstract Kleptoparasitism is a specialised form of foraging interference occurring throughout the animal kingdom and especially is well documented among birds including African fish eagle Haliaeetus vocifer. However, only one record is reported between fish eagle and shoebill Balaeniceps rex despite their habitat overlap and it remains undescribed. This paper documents kleptoparasitism of shoebills by fish eagles in Malagarasi wetlands, western Tanzania. Kleptoparasitism was highly seasonal occurring in the dry season, especially at low floods. Both adults and non-adult eagles stole prey from shoebills, with non-adults initiating attacks at shorter distances than adults perhaps because of the lack of experience. The shoebill preys were comparatively larger than those from fish eagle own fishing and required long preparation time which provided the opportunities to kleptoparasites. Shoebill’s long prey handling time (6.90 ± 4.48 min) may have provided benefits to fish eagles as all kleptoparasitism attempts (n = 138) were successful. Moreover, it seemed likely that an attacking fish eagle posed significant danger to the shoebill and given the fact that shoebill lacks structures such as pointed bill to defend from kleptoparasitic attack it opted not to retaliate. Perch availability and habitat openness in the study sites could also have facilitated kleptoparasitic behaviours. Keywords: Balaeniceps rex, Haliaeetus vocifer, handling time, kleptoparasitism, Malagarasi- Muyovozi Ramsar Site Introduction openness (Paulson 1985), large concentrations Kleptoparasitism is a specialised form of of hosts (Brockmann and Barnard 1979), foraging interference (Perrins and Birkhead large/visible food items or prey (Brockmann 1983) occurring throughout the animal and Barnard 1979, Dekker et al. 2012) and kingdom (Iyengar 2008), and especially is shortage of food from self-foraging (Oro well documented among birds (Brockmann 1996). Moreover, age of kleptoparasites and and Barnard 1979). It is particularly common hosts as well as the distance between them; in waterbirds (Furness 1987) and in have also been hypothesized to play Accipitridae (Morand-Ferron et al. 2007) significant roles in avian kleptoparasitism especially eagles; notably the African fish (Broom and Ruxton 1998, Dekker et al. 2012). eagle Haliaeetus vocifer (Sumba 1989, Distances between the hosts and Kasoma 1995) and bald eagles Haliaeetus kleptoparasites not only affect the chances of leucocephalus (Dekker et al. 2012). observing kleptoparasitic opportunities but Several studies have suggested ecological also increases the cost of attempting conditions that relate to the evolution of kleptoparasitism as kleptoparasites expend kleptoparasitism; they include habitat energy when travelling to the hosts. This 131 http://journals.udsm.ac.tz/index.php/tjs www.ajol.info/index.php/tjs/ John and Lee- Kleptoparasitism of shoebills by African fish eagles in Western Tanzania could be more important for raptorial Nahonyo and Msuya (2008) registered one kleptoparasites such as fish eagles that perch incidence of kleptoparasitism on a shoebill by on top of trees or emergent vegetation fish eagles in Malagarasi wetlands but the overlooking hosts. interaction remains undescribed. Specifically, Kleptoparasitised individuals may in addition to documenting this challenge or retaliate, escape with food to kleptoparasitic interaction, this research cover, reduce handling time (including hypothesized that; (1) the rate of switching to smaller prey) or remain passive kleptoparasitic attack would be higher in (Stillman et al. 1997). Stillman et al. (1997) driest months when fish eagles may be hypothesized that kleptoparasitism should be precluded from catching prey and it is during flexible, with aggression only occurring when this period when shoebills become restricted benefits of this action outweigh the costs. to fewer and wetter locations which may Sirot (2000) suggested that kleptoparasitism attract kleptoparasites, and (2) the prey size and aggressiveness should increase when food and handling time would differ between hosts patches are harder to locate. Using ideal free foraging under pressure of kleptoparasites and distribution (IFD) model, Hamilton (2002) those that foraged in areas without found that kleptoparasitism increases with kleptoparasites. increasing difference in searching efficiency, increasing difference in fighting ability, Materials and Methods increasing handling time while decreases with Study sites increasing resource input rates, increasing Field work was conducted between June ownership advantage and increasing fighting 2011 and February 2012 within the central time. In general, when payoff for drainage of the Malagarasi-Muyovozi Ramsar kleptoparasitism is low, individuals switch to Site (3–6°S, 30–32°E), and largely within a searching for prey (Stillman et al. 1997, Smith swamp located in southern part of Lake et al. 2002). Nyamagoma around the village of Kasisi This paper both qualitatively and (5°03'05"S, 30°57'14"E) (Figure 1). The quantitatively describes kleptoparasitic water levels in the lake and adjacent interactions between shoebills Balaeniceps floodplains can fluctuate widely on an annual rex and fish eagles. The two are large basis depending on rainfall and inflow waterbirds in sub-Saharan Africa where in (Nkotagu and Ndaro 2004); annual flooding some large wetlands (e.g., in Sudan, Tanzania, beginning in November and peaking between Uganda and Zambia) they share habitats April and May. Large and small open water (Brown et al. 1982). However, despite the channels around this lake variously open and habitat overlap between the two species, there close depending on currents and water volume. is no mention of kleptoparasitism in The vegetative swamp surrounding the edges Balaenicipitidae family (Gould 1851, Hagey of the lake is largely comprised of grasses and et al. 2002), of which shoebill is the only sedge species. member, in published reviews (Brockmann At Kasisi village, the miombo and Barnard 1979, Morand-Ferron et al. 2007, Brachystegia woodland which is mixed with Iyengar 2008) and in shoebill foraging studies Borassus palm (Borassus aethiopum) (Figure in Sudan, Uganda and Zambia (Guillet 1979, 2a, b) bordering the swamp on one side. In Möller 1982, Mullers and Amar 2015). The between the tall vegetative swamp and most recent extensive review of woodland, there are extensive open glades of kleptoparasitism in birds being that of short vegetation dominated by Eleocharis- Morand-Ferron et al. (2007) where 33 Leersia plant species forming dense floating families were noted to be involved in vegetation platforms (John and Lee 2012). interspecific kleptoparasitism. Moreover, These floating platforms sometimes contain 132 Tanz. J. Sci. Vol. 45(2), 2019 small natural surface water openings (pools), by shoebills to hunt for air-breathing fish that which are supplemented by those made by become sparse, especially as water recedes. humans and the antelope Sitatunga (Tragelaphus spekii). These ‘pools’ are used Figure 1: Map showing the study area at Kasisi Village, South of Lake Nyamagoma, within Malagarasi-Muyovozi Ramsar Site. Study species eastern sub-Saharan Africa from Sudan to The African fish eagle is a widespread Zambia (Feduccia 1977, Hancock et al. 1992) raptor in Africa occurring at many water and therefore classified as ‘vulnerable’ by bodies containing fish except in arid areas IUCN (BirdLife International 2019). Most of (Brown et al. 1982). As its name suggests, it the shoebill habitats are treeless remote chiefly feeds on fish, although other prey such floodplains and permanent swamps (Hancock as rodents, waterbirds and reptiles are et al. 1992). The shoebill is a rare (e.g., < 500 occasionally consumed (Stewart et al. 1997). mature individuals survive in Tanzania) It is, however, the most known kleptoparasitic wetland specialist large bird (John et al. 2012). raptor in Africa (Sumba 1989, Kasoma 1995). The shoebill has extremely long toes (Gould The shoebill has a narrow and disconnected 1851) to support its weight (≈ 7 kg) when distribution along major water basins in walking on floating vegetation platform 133 John and Lee- Kleptoparasitism of shoebills by African fish eagles in Western Tanzania unlike other large wetland birds which have to ambushing of surfacing prey, a phenomenon wade. The foraging behaviour of the shoebill that Guillet (1979) described as ‘collapsing’ resembles that of many Ardeidae, i.e., ‘stand- (Figure 2c). The shoebill is a non-social and-wait’ and ‘walk slowly’ (Guillet 1979, forager (Möller 1982) but may form loose Mullers and Amar 2015), but in shoebill this groups at good foraging sites. is usually followed by a pronounced (a) (b) (c) (d) Figure 2: (a) An adult African fish eagle (Haliaeetus vocifer) perched on tall sedge clump attending the shoebill (Balaeniceps rex) for food stealing,