@ @ B D 9+82 ;7 +*8 doi: 8+ 87788E 9+82 +*8 http://folia.paru.cas.cz
Research Article Feeding strategies and competition between terrestrial Haemadipsa leeches (Euhirudinea: Arhynchobdellida) in Danum Valley rainforest (Borneo, Sabah)
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F ! ! $ G H! $ $ G ! G ! ! % ! [ J - ography and taxonomy. We undertook research on two species inhabiting lowland dipterocarp forest, Haemadipsa picta K 8'9' and Haemadipsa subagilis MK 8'9'N ! & P M N ! ƽ ! G G & ƽ- R= MGN G ! J - [ R= M N - [ G ƽ G ! $ $ ! R S ! & ! H. picta is more G ! ƽ- $ & ! ! = ! J & ! plant height. Haemadipsa subagilis & U! G ! V G & [ G & ! M N J - H. picta M ! & ! [ N= (ii) habitat specialisation of H. subagilis. Moreover, we provide new observations on their foraging behaviour. ectoparasites, foraging behaviour, Haemadipsidae, haematophagy, parasitism
H B !-> ! G $ - $ [ ! & $ ! - ! ! F - G ! ƽ M> ! ! ! G N &! ! ! G P B - $ 1+Z ! ! G ! [ $ ! ! - ! Haemadipsa H 81<' M brown leech species complex (former H. zeylanica sensu B 9++7 9+8+ B H % 9++1N H! ! lato B 9+8+N P - fauna of the Oriental region has received much attention mary tropical rainforest, whereas more disturbed environ- ! & K M8'9' 8'*< 8'*1N ments are dominated by conspicuously coloured H. picta. The tiger leech H. picta K 8'9' & G & ! - coloured species, namely H. subagilis MK 8'9'N gies of hematophagous leeches, have not been examined H. sumatrana MF 811*N & - G \ ! ! & mortality and, therefore, their abundance in an ecosystem MB & 8'19 $ ! 9++7N M K 8'21N F & $ G Many ecological aspects of the life history of haemad- simply apply this theory to ectoparasites, such as leeches, ipsid species remain unresolved, including establishing &! ! ! ! B an average life span for each species and species habitat G G & ! U M D M8'1 ! " # $ % & ' *+-*12 3 & ! 671 89;;7;7<<= >- ? This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. 8+ 87788E 9+82 +*8 b > G ! $ Haemadipsa leeches AB S1 S2 P1 P2 P3 P4 P5 2 km 50 km Fig. 1. A M j [ _ @ $ M[_@ N B G ! MK N= B M [ _ D @ M[_D@N $ ! M 8M D 9+82 ;7 +*8 9 2 8+ 87788E 9+82 +*8 b > G ! $ Haemadipsa leeches A 24 B 7 22 6 20 18 5 16 14 4 12 10 3 8 2 6 4 1 2 0 0 Fig. 2. $ G $ M 8++ 9 plot) occurring in the particular type of habitat. A – Haemadipsa picta K 8'9'= B – H. subagilis MK 8'9'N H! ' 31 H& -& `S_ $ [ - 30 MD { 2 81 { + +9N ! G H. pic- 9 29 ta G & M $ G $ E8++ : 28 `t { 8; < / 8 'N ƽ- M`o { ' 9 / 8 1N ! % ! ! 27 ` ! G H. subagilis were found 26 MD { 9 87 { + 82= `t { 9 ; / + < `o { 7 + / 8 += 25 D 9 G ! N ! 24 - [ M { + 21 + 91 $ N 23 S -& `S_ [ ƽ $ 22 body length (L) of H. picta collected during the morning Mj { 91 + / 8 +N Mj { 9* ; / 8 *= 21 D { ; 1' { + +8= D *N & ! ! M 20 ! ! J & H G 8N ! ! 3 -V $ - ferences in the case of H. subagilis MF { + <8 { + 71N Fig. 3. An average body length (mm) of Haemadipsa picta K 8'9' $ ! MKN - [ $ G & M N H! ' D 9+82 ;7 +*8 * 2 8+ 87788E 9+82 +*8 b > G ! $ Haemadipsa leeches 50 A 40 30 20 10 B 0 0 20 40 60 80 100 120 140 160 180 200 220 Fig. 4. The dispersion plot depicting the correlation between the length and preferred altitude of attack of Haemadipsa pic- ta K 8'9' H! G -[ ! $ & ! & P ! $ { *+ ;19 & 8+MUN - 99 29<9 The present study shows that H. picta exhibits oppor- tunistic behaviour and agglomerates in the vicinity of ani- mal trails used mainly by mammals such as humans, beard- P &! ! Fig. 5. Typical postures of haemadipsid leeches on the example of increase in hunt success ratio. The average number of col- Haemadipsa picta K 8'9' A M = B – in at- tack position. Photo © J j & & ! $ & $ ! ! ƽ- ! G is worth mentioning that host preferences for both species & G B ! M9+8 D 9+82 ;7 +*8 7 2 8+ 87788E 9+82 +*8 b > G ! $ Haemadipsa leeches ! # [ $# # ) QG & $ R U G ! $ MD < N ! [ Haemadipsa typically found in the larger individuals (adult), and the lat- B ! ! G ! $ G % $ Haemadipsa picta terrestrial genera whose members very probably do not is generally more active and moves more rapidly than its compete for food resources: many species of Haemadip- smaller congener, and very often chases potential hosts. sa G = ! Tritetrabdella that B G ! $ & G $ H. subagilis. ! G M ! ` B M9N H. subagilis attacks from ambush, i.e. it spends 9+87N= G Phytobdella ! 81'7 ! a considerable amount of time in a resting posture, and be- MK 8'*1N H! ! Planob- comes active and attaches to a host when it comes very della ! 81'7 & M ! G ! ! 817'N H! ! G & $ - individual almost always performs head movements, but cies, such as of H. picta and H. subagilis, were hitherto un- rarely demonstrates body waving. ravelled. Given that both species occur in lowland forests M*N V! H. subagilis ! ! ! R G - in very high numbers, one would ask how it is possible most never crawls on its host, instead it tries to penetrate ! - U & ! ! ! the skin near the site of initial attachment. Our survey suggests that H. picta reduces the level of M7N [ ƽ G & $ & [ G $ - [ G $ B & cal distribution of individuals, thereby utilising a range of M8'18N ! & ! & !- ƽ ! D ! [ & $ ƽ G ! ! ! $ & ƽ & [ body loop during movement, as exhibited by H. ghilianii P ! $ G G $ ! ! G G J H. pic- & ! ! ! % $ ! ! M ta !& & % $ G $ - example, rodents), and adults to larger hosts (pigs and pri- form crawling once an individual matures. Haemadipsa N= ! ! ! subagilis, although capable of moving in these ways, was G & J > - observed to prefer vermiform crawling, irrespective of age. M9+8;N ! & ! ! G J The repertoire of foraging behaviours of other cryptical- with tick diversity, namely, the larger the host, the more ly coloured species of Haemadipsa presumably resembles $ ! ! G ! that of H. subagilis. Our observations illustrate the behav- host body size also determines the body size in land leech- G ! ! - es. the uniform environment of the dipterocarp rainforest. Although we did not show how H. subagilis ƽ ! $ [ ! J - , ), , & [ & & [ Haemadipsa picta aggregates along trails in order to in- over half of collected individuals came from riparian local- G G [ ! &! ! ! - ities. This suggests that H. subagilis occupies a more spe- & [ H. subagilis. Furthermore, cialised niche in comparison with H. picta, presumably due G ! J - H. picta, whereas in to being more sensitive to low humidity (the best evidence H. subagilis, which prefers ambush in the leaf litter, it ap- for this is the overall number of H. subagilis vs H. picta G - [ D H. picta ƽ obtained during the dry season). Overall, our results are ! [ G G $ ! & ! ! G G " ! $ \ [ G F G M9++;N & ! G ! separation of feeding in subadult and adult stages. Further same environment because of very likely asymmetrical studies are needed to discover habitat preferences of H. sub- antagonism between them. This results in attenuation of agilis F ! ! G $ ƽ $ $ $ ƽ & ! ! ! ! Haemadipsa leech- G G G ! \ ! G G & - es, H. picta appears to be the stronger competitor, shaping asite and host populations system, making the coexistence the ecological relationship between species. G ! - [` M B ! 9+8 D 9+82 ;7 +*8 < 2 8+ 87788E 9+82 +*8 b > G ! $ Haemadipsa leeches + $ Authors are very grateful to two anon- [ D J % V & K % J ymous reviewers for their remarks and linguistic corrections, for explorative discussions on various aspects of their research. &! ! $ ! - H! & ! H > D cal reading. Moreover, authors like to thank to their course tutor, @ ! " # $ & G ! K @J B M > $ B " - > $ B D # $ N $ > ! B M [BEV `S E `SBE2<2E9+8;N H! J j & M > $ B " - authors declare that they have no competing interests. # $ N [ ! ! REFERENCES ǏDždžǓǔǐǏ K Kǂǚ K 8'21 G 3ǖǕǔDŽljdžǓǂ # 9++* H! ! ! Erpob- ! - " della octoculata Mj N G $ F G 11 > 72 98'M972 '7M8+8 ǂǔljdžǚ D BǂǓNJǏ H jNJǗdžǍǚ @ K 9+8; - jǂNJ -H @ljdžǏ " -F 9+8+ j ! 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