Food Habi s o Cynop e us b achyo is (Mulle ) (Chi op e a: P e opodidae) in Peninsula
Malaysia
Au ho (s): K. H. Tan, A. Zubaid and T. H. Kunz
Sou ce:
Jou nal o T opical Ecology
, May, 1998, Vol. 14, No. 3 (May, 1998), pp. 299-307
Published by: Camb idge Uni e si y P ess
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Jou nal o T opical Ecology (1998) 14:299-307. Wi h 1 igu e
Copy igh ? 1998 Camb idge Uni e si y P ess
Food habi s o Cynop e us b achyo is (Mulle )
(Chi op e a: P e opodidae) in Peninsula
Malaysia
K. H. TAN*, A. ZUBAID*' and T. H. KUNZ
*Depa men o Zoology, Uni e si i Kebangsaan Malaysia, 43600 UKA Bangi, Selango ,
Malaysia (e-mail: [email p o ec ed])
Depa men o Biology, Bos on Uni e si y, Bos on, Massachuse s 02215, USA
(Accep ed 26 No embe 1997)
ABSTRACT. In o ma ion on he eeding habi s o he lesse dog- aced ui ba ,
Cynop e us b achyo is, was ob ained by he collec ion o ood emains di ec ly benea h
day ime and eeding oos s. The ba s we e ound o eed on he ui s o 54 plan
species, he lea es o 14 species and he lowe pa s o ou species. The seasonal
phenological di e ences among congene ic plan species led o a s eady p oduc ion
o ui h oughou he yea and he da a sugges ha Ficus spp. a e a key compon-
en in he die . Judging om i s wide selec ion o ui s, C. b achyo is, is conside ed
o be an impo an seed dispe se . Foli o y in C. b achyo is appea s o be mo e
common han p e iously hough . O he lea es consumed by he ba s, se en spe-
cies belonged o he amily Leguminosae, ollowed by My aceae, Mo aceae, Rhizo-
pho aceae and Eupho biaceae. F ui s, in gene al, p o ide an ene gy- ich die o
phy ophagous ba s bu mos a e low in p o ein. In con as , lea es consumed by
ba s ha e a ela i ely high p o ein con en . We sugges ha oli o y (by lea
ac iona ion) should be ene ge ically mo e ad an ageous han he inges ion o
la ge amoun s o low p o ein ui o he ac i e pu sui o mobile insec s.
KEY WORDS: Cynop e us b achyo is, Ficus, lowe s, oli o y, ui s, Malaysia, second-
a y o es
INTRODUCTION
All species o p e opodids a e hough o be mainly phy ophagous, aking ui ,
lo al esou ces (nec a and pollen), o lea es (Fleming 1982, Ki chene e al.
1990, Kunz 1996, Kunz & Diaz 1995, Lim 1970; Ma shall 1983, 1985; Phua &
Co le 1989) wi h insec s o ming a small componen (Funakoshi e al. 1993,
'Co esponding au ho .
299
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300 K. H. TAN, A. ZUBAID AND T. H. KUNZ
Lim 1970). In he wild, he eeding a eas and he composi ion o hei ood a e
la gely in luenced by he seasonal lowe ing and ui ing o ees (Lim 1966).
Ma shall (1983) sugges ed ha phy ophagous ba s a e nei he ue gene alis s
no specialis s, bu a he 'sequen ial specialis s', a ou ing a any ime one o
a ew plan species amongs po en ial ood plan s a ailable a ha season.
The lesse dog- aced ui ba , Cynop e us b achyo is (Mulle ), is a common
ugi o ous species in Sou heas Asia. Th oughou i s ange, his ba occupies
a a ie y o habi a s including p ima y o es , dis u bed o es , mang o e, cul-
i a ed a eas, o cha ds, ga dens and u ban a eas (F ancis 1990, 1994; Lim
1966, Medway 1983, Phua & Co le 1989; Zubaid 1993, 1994). This species is
well adap ed o an h opogenic condi ions. In an ea lie s udy (Tan e al. 1997),
we showed ha C. b achyo is modi ied he lea es o palm ees o cons uc en s
which we e han used as day and/o eeding oos s. Apa om iden i ying
sui able oos ing si es, i is also impo an o know he kinds o ood esou ces
ha sus ain hese popula ions in u ban a eas. He e we epo some p elimina y
da a on he ood habi s o C. b achyo is in an u ban en i onmen .
METHODS
Th ee s udy si es we e selec ed based upon su eys o e idence o palm en s
and en - oos ing ba s (Tan e al. 1997). Only C. b achyo is was ound o con-
s uc and u ilize hese en s. These si es we e loca ed on he campus o Uni -
e si i Kebangsaan Malaysia (UKM) in Bangi, Uni e si i Pe anian Malaysia
(UPM) in Se dang, and Zoo Nega a (Na ional Zoological Pa k) in Hulu Kelang.
All o he s udy si es a e ex ensi ely landscaped wi h ees and can be consid-
e ed as seconda y o es habi a s. In addi ion, he eas e n and sou he n sides o
he UKM campus a e bo de ed by he Bangi Fo es Rese e (dis u bed lowland
dip e oca p o es ), c. 100 ha in size. Desc ip ions o he lo is ic composi ion
o his o es a e gi en in La i (1990).
Obse a ions we e made om Janua y o Decembe 1996. In o ma ion on
he eeding habi s o Cynop e us b achyo is we e ob ained by collec ing ui em-
nan s, seeds, lea es and lowe pa s d opped di ec ly benea h eeding and
day ime oos s. Collec ion o ood samples we e ca ied ou a leas once each
week. The u iliza ion and a ailabili y o ui s, lowe s and lea es we e de e -
mined on a 'p esence/absence' basis (Thomas 1988). All he plan species we e
iden i ied using Co ne (1988), K empin (1993), Ng (1978, 1989) and Whi -
mo e (1972, 1973), and we e u he e i ied using he he ba ia a UKM and
he Fo es Resea ch Ins i u e o Malaysia (FRIM) a Kepong.
RESULTS
Cynop e us b achyo is was obse ed o eed upon he ui s o 54 plan species,
he lea es o 14 species and he s amens o ou species (Tables 1 and 2). The
ui bea ing amilies ha domina ed he lis a e: Sapo aceae (nine species),
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Food habi s o Cynop e us b achyo is 301
Table 1. F ui emnan s collec ed benea h eeding and day- ime oos s o Cynop e us b achyo is.
Plan species Family* Loca ion Jan Feb Ma Ap May Jun Jul Aug Sep Oc No Dec
Pipe aducum 1 a + + + + + + + + + + + +
Te minalia ca appa 2 ab + + + + + + + + + + + +
Mun inzgia calabu a 3 ab + + + + + + + + + + + +
Calophyllum inophyllum 4 abc + + + + + + + + + + + +
Ficus is ulosa 5 ac + + + + + + + + + + + +
Elaeoca pus s ipula is 3 c + + + + + + +
Eugenia g andis 6 abc + + + + + +
Elaeoca pus sp. 3 c + + + +
Eugenia sp. (2) 6 c + + + +
Musa sp. 7 a + + + + + +
F ag ae ag ans 9 b + + + + + +
Ach as zapo a 8 b + + + + +
Psidium guaja a 6 ab + + + +
Eugenia aquea 6 abc + + + + + + + +
Polyal hia longi olia 10 ac + + + + + + +
Pellacalyx sacca dianus 11 c + + + + + + +
Ficus oxbu gii 5 c + + + +
Flacou ia ine mnis 12 b + + +
Mimusops elengi 8 bc + + + + + + +
Ficus sp. (1) 5 a + +
Eugenia malaccensis 6 c + + + + +
Li is ona o undi olia 14 ac + + + + +
Nepheliumn malaccensis 13 c + + +
Licuala g andis 14 c + + + +
G ewia omnen osa 16 a + + +
P e nand a echina a 15 d +
Ficus benjainina 5 ac + + +
Mangi e a indica 10 b + +
Ficus eligiosa 5 b + +
Ch ysalindoca pus lu escens 14 c + +
Li is ona chinensis 14 bc + +
P ycospe ma maca hu ii 14 c +
Annona squamnosa 10 b + +
Roys onea egia 14 c + +
Eugenia jambos 6 bc + + +
A oca pus maingayi 5 c + + +
Palaquium obo a uni 8 c + + +
Payena maingayi 8 c + + +
Pou e ia malaccensis 8 c + + +
Eugenia sp. (1) 6 c + +
Diospy os sp. 17 c + +
Palaquium gu a 8 c + +
Palaquiun cla keanunm 8 c + +
Payena lucida 8 c + +
Madhuca selango ica 8 c + +
Ficus sp. (2) 5 c + +
Ficus sp. (3) 5 c + +
Eugeniia sp. (3) 6 c +
Uniden i ied seeds (1) ? a + + +
(2) ? a + +
(3) ? c + +
(4) ? c +
(5) ? c + +
(6) ? c +
* 1 = Pipe aceae, 2 = Comb e aceae, 3 = Elaeoca paceae, 4 = Gu i e ae, 5 = Mo aceae, 6 = My aceae, 7 =
Musaceae, 8 = Sapo aceae, 9 = Loganiaceae, 10 = Anaca diaceae, 11 = Rhizopho aceae, 12 = Flacou iaceae,
13 = Sapindaceae, 14 = Palmae, 15 = Melas omaceae, 16 = Tiliceae, 17 = Ebenaceae
a = Zoo Nega a, b = Uni e si i Pe anian Malaysia, c = Uni e si i Kebangsaan Malaysia
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302 K. H. TAN, A. ZUBAID AND T. H. KUNZ
Table 2. Chewed lea es and lowe s collec ed benea h eeding and day- ime oos s o Cynop e us b achyo is.
Food
Plan species Family* Loca ion ype Jan Feb Ma Ap May Jun Jul Aug Sep Oc No Dec
E y h ina
o ien alis 1 a Lea + + + + + + + + + + + +
E y h ina
subumb ans 1 a Lea + + + + + + + + +
Eugenia g andis 2 bc Lea + + + + + +
A oca pus
ul ico ex 3 c Lea + + + + + + + + +
Cassia spec abilis 1 b Lea + + + + + + +
Cassia spec abilis 1 b Flowe + + + + + + +
Pel opho um
p e oca pum 1 c Flowe + + + + + +
Ficus eligiosa 3 b Lea + + +
Pellacalyx
sacca dianus 4 c Lea + + + + + +
Cassia is ula 1 b Lea + + + +
Cassia is ula 1 b Flowe + + + +
Bauhinia
pu pu ea 1 c Flowe + + +
E y h ina sp. 1 b Lea + + +
E y h ina glauca 1 c Lea + +
Eugenia aquea 2 c Lea + +
E y h ina
a iega a 1 bc Lea + + +
He ea b asiliensis 5 c Lea + + +
Eugenia sp. (1) 2 c Lea + +
* 1 = Leguminosae, 2 = My aceae, 3 = Mo aceae, 4 = Rhizopho aceae, 5 = Eupho biaceae
a = Zoo Nega a, b = UPM, c = UKM (as Table 1)
Mo aceae (eigh species), My aceae (eigh species), and Palmae (six species).
O he lea es consumed by he ba s, se en species belonged o he amily Legu-
minosae. This was ollowed by My aceae, Mo aceae, Rhizopho aceae and
Eupho biaceae. All he lowe s belonged o he amily Leguminosae.
F ui s o i e species we e ound h oughou he yea , namely Te minalia
ca appa, Mun ingia calabu a, Pipe aduncum, Calophyllum inophyllum and Ficus is u-
losa (Table 1). O he plan species ha had a long ui ing season we e Elaeo-
ca pus s ipula is, Eugenia g andis and Polyal hia longi olia. Al hough ui s o he
abo e-men ioned species (excep Mun ingia calabu a) a e ound in abundance
in he UKM campus, C. b achyo is he e p e e ed o eed on a ious kinds o
ui s om he Bangi Fo es Rese e du ing he ui ing season ( om July o
Sep embe ) (Table 1). In he Bangi Fo es Rese e, mos o he ui ing ees
we e c. 20 m in heigh . These included Palaquium spp, Payena spp., and Pou e ia
malaccensis. O namen al palms p o ide an impo an ood sou ce o C. b achyo is
in he mon hs o June, ea ly July and Oc obe , jus be o e and a e he o es
ui ing season. So ui s we e ei he o ally o pa ially consumed. Howe e ,
i was qui e common o some ui s o be d opped undamaged unde he
eeding oos s o day- ime oos s. In he Bangi Fo es Rese e, unea en ui s
wi h ba oo h ma ks we e o en obse ed on he o es loo .
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Food habi s o Cynop e us b achyo is 303
The lea es and lowe s upon which C. b achyo is eeds a e a ailable h ough-
ou he yea and hei consump ion was non-seasonal (Table 2). Cynop e us b a-
chyo is appea ed o eed ex ensi ely on he lea es o Eiy h ina o ien alis, E. subum-
b ans, Eugenia g andis, Cassia spec abilis, Pellacalyx sacca dianus and A oca pus
ul ico ex. In he case o he Eugenia spp., A oca pus ul ico ex and Ficus eligiosa,
only he young lea es we e ea en. The lowe s o Pel opho um p e oca pum, Bauhi-
nia pu pu ea, Cassia spec abilis and Cassia is ula we e equen ly isi ed by C.
b achyo is.
All o he lea pelle s we e collec ed unde he day ime oos s and eeding
oos s. The lea es we e chewed and he soluble con en s ex ac ed by he ba s.
The ib ous con en s we e disca ded as ib ous pelle s. Pe ioles and eins we e
common in he lea pelle s. Di e en plan species may p oduce di e en ly
shaped lea pelle s. The lea pelle s o Egy h ina o ien alis we e usually 'kidney'
o 'm' in shape (Figu e 1), wi h he a e age leng h and wid h measu emen s
(n = 60) o 1.18 ? 0.11 cm and 0.70 ? 0.23 cm, espec i ely. On he o he hand,
he lea pelle s o A oca pus ul ico ex we e mos ly ound o o oid in shape,
1.08 0.17 cm and 0.84 ? 0.15 (n =60).
i l l | |~~~~~~~~~~~.. . ....... R ,.$ o..:<..'';..../g
~~~~~~~~~~~~~~~~~~~~~~~~~~~... . . .. .. .. . ..... ' ; ..
and da im os o cyno e s b$ achyo zs> ;X
. . . . .. .;. .. . i--@e&>g'
e. . . '' '':' ... ... . .
... j R.. ......... 2. ..... . ,.... ..
<. . l ..^.'-'. kS'lb i e' "' ; s''.... .....
ij . l ~~~~. .. .... |. . : i. . e i > ixj
Fiu . P ill chee le e an eece lea pele o Eh n o e ai cole e bena eing
DISCUSSION
F ui s ha a e a ailable h oughou he yea o wi h a long ui ing season a e
p esumably a majo ac o in main aining he popula ion o C b achyo is. In
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304 K. H. TAN, A. ZUBAID AND T. H. KUNZ
addi ion, he seasonal phenological di e ences among congene ic plan species
(e.g. Ficus spp. and Eugenia spp.) led o a s eady p oduc ion o ui h oughou
he yea . The p esen da a indica es ha Ficus spp. may be an impo an com-
ponen in he die s o C. b achyo is. P e ious s udies ha e shown ha p e opodid
ba s ha e a g ea p edilec ion o Ficus spp. (Bha 1994, Ma shall 1985, Phua &
Co le 1989). Simila ly, some phyllos omid ba s a e also known o o age la g-
ely on Ficus spp. (Fleming & Hei haus 1981, Handley e al. 1991; Mo ison 1978,
1980). Du ing he o es ui ing season, C. b achyo is p e e ed o o age in he
o es a he han in u ban a eas. This indica es ha C. b achyo is shows dis inc
ood p e e ences a any one place and ime i a choice is a ailable. Mos o he
ba ui s in he o es we e loca ed a he canopy le el which may explain why
p e opodid ba s a e mo e eadily cap u ed he e han in he unde s o ey
(F ancis 1990, 1994; Zubaid 1994).
In his s udy, C. b achyo is was obse ed o isi lowe s o ou legume species.
Lim (1970) and Ki chene e al. 1990 ound conside able quan i ies o pollen
in he aecal samples o C. b achyo is. Phyllos omid ba s ha e been obse ed
eeding on he lowe s o Bauhinia spp. (Fische 1992, Hei haus e al. 1974). I
is gene ally ag eed ha ba s isi lowe s o nec a , pollen and lo al pa s
(Bake & Ha is 1957, Ma shall 1985). Rasweile (1977) sugges ed ha pollen
may p o ide an impo an sou ce o die a y p o ein o plan - isi ing ba s.
In addi ion o ui s and lowe s, C. b achyo is also ed egula ly on he lea es
o 14 plan species (Table 2) and is he i s published epo o he impo ance
o his componen o his species. Lim (1970) s a ed ha lea es ep esen ed
a mino componen o he die whe eas Ki chene e al. (1990) no ed he p es-
ence o plan epide mis in hei analyses. Phua & Co le (1989) who collec ed
plan emains unde eeding oos s only epo ed he p esence o ui s and
seeds. O hese, he mos impo an we e E j h ina spp., Cassia spp., Eugenia
spp. andA oca pus ul ico ex. Bha (1994) epo ed ha C. sphinx ed on lea es
o Cassiajis ula and E j h ina indica. E y h ina spp. a e also commonly ea en by
o he phy ophagous ba s (Kunz & Diaz 1995, Ma shall 1985, Zo ea & Mendes
1993).
Se e al hypo heses ha e been p oposed o explain why phy ophagous ba s
eed on lea es. F ui s a e ypically high in ca bohyd a es and poo in p o ein
(Rasweile 1977, Thomas 1984a). Phyllos omid ba s which ha e he abili y o
echoloca e can consume insec s in o de o pa ially ul il hei p o ein equi e-
men s. Unlike phyllos omid ba s, p e opodids do no echoloca e, hence, i is
e y di icul o hem o hun o insec s. Thomas (1984) and S ella (1986)
sugges ed ha p e opodids may o e -consume and hen exc e e ca bohyd a es
in o de o mee hei p o ein equi emen s om o he wise p o ein-poo ui s.
Al e na i ely, lea es could p o ide an impo an sou ce o die a y p o ein o
phy ophagous ba s (Kunz 1996, Kunz & Diaz 1995, Kunz & Ingalls 1994, Low y
1989, Rasweile 1977, Thomas 1984a). Low y (1989) epo ed ha P e opus elec o
ed ex ensi ely on Albizia lebbek lea es (36% p o ein on d y ma e basis) by
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Food habi s o Cynop e us b achyo is 305
ex ac ing he p o ein- ich soluble ac ion and ejec ing he p o ein-poo ib-
ous pelle s. Simila ly, A ibeus jamaicensis is known o consume lea es o
E j h ina poeppigiana which a e ela i ely high in p o ein (19% d y ma e
con en ) (Kunz & Diaz 1995). Besides ba s, Choo e al. (1981) no ed ha
E j h ina excelsa (17% p o ein on d y ma e basis) se ed as an impo an p o-
ein sou ce o some neo opical p ima es. In his s udy, lea es o E j h ina spp.
and Cassia spp. we e p e e ed by C. b achyo is. The p o ein con en o E j h ina
a iega a and Cassia spec abilis has been es ima ed as 21.7 and 30.7% o d y
ma e , espec i ely (Telek & Ma in 1983). Some phyllos omid ba s may
p e e o consume p o ein- ich lea es (Kunz & Diaz 1995, Zo ea & Mendes
1993), because lea es a e abundan , and p edic able in space and ime (Kunz &
Diaz 1995). Ba s need less ene gy o pu sue lea es as compa ed o insec s.
Hence, ba s can educe hei daily ene gy expendi u e especially du ing la e
p egnancy and lac a ion when ene gy and nu ien demands a e expec ed o
be highes . Kunz & Diaz (1995) concluded ha he consump ion o lea es by
ba s p o ides a highe yield o p o ein pe uni o aging e o .
The lea es could also be an impo an sou ce o mine als and i amins.
Lea es o some plan species a e ich in calcium and phospho us as compa ed
o ui s (He e a 1987, Rasweile 1977). Rasweile (1977) epo ed a high
mo ali y o cap i e p e opodids main ained o ex ended pe iods on ui s due
o calcium de iciency. Females bea almos he en i e nu ien cos o p odu-
cing he skele on o hei o sp ing (Ba clay 1994, Papadimi iou e al. 1996).
Recen e idence sugges s ha he e is a high demand o calcium and a sca -
ci y causes signi ican s uc u al changes in he bones o lac a ing ba s
(Kwiecinski 1987 e al.).
Lea es may also con ain se e al me aboli es impo an o ep oduc ion in
ba s (Cunningham an Some an 1972, Kunz & Diaz 1995). These me aboli es
include s e oidal sapogenins and alkaloids ( e iewed by Kunz & Diaz 1995).
Since s e oids a e essen ial o ho mone syn hesis, he o al in ake o hese
compounds by phy ophagous ba s may in luence hei ep oduc i e ac i i y
(Wickle & Seib 1976). On he o he hand, i is well documen ed ha second-
a y me aboli es especially phenolic compounds (e.g. condensed annins) can
ac as inhibi o s o p o ein diges ion (Telek & Ma in 1983). Since p e opodids
ha e sho diges i e ac s and apid ansi - imes (Fo man 1990, Richa dson
e al. 1987, Tedman & Hall 1985), a high concen a ion o phenolic compounds
could limi he e ec i eness o p o ein diges ion. Howe e , lea es consumed
by ba s, such as E y h ina spp. and Albizia spp. a e ela i ely low in condensed
annins and o al phenols (Low y 1989, Telek & Ma in 1983). Mo eo e , young
lea es con ain highe p o ein con en and lowe seconda y me aboli es han
ma u e lea es (Telek & Ma in 1983). The la e may explain why C. b achyo is
p e e s o consume young lea es o Eugenia spp., Ficus eligiosa and A oca pus
ul ico ex.
We sugges ha oli o y (by lea ac iona ion) may e lec high p o ein and
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306 K. H. TAN, A. ZUBAID AND T. H. KUNZ
low phenolic con en o lea es. In gene al, ui s p o ide an ene gy sou ce
(ca bohyd a es) o phy ophagous ba s, and lea es may be an impo an sou ce
o p o ein, mine als, and s e oids. Pollen is also a p o ein- ich ood and pe haps
lea es a e consumed o a g ea e ex en when pollen is una ailable. Fu u e
s udies should in es iga e he nu i ional composi ion o he ui s, lea es and
lowe s ea en by hese ba s.
ACKNOWLEDGEMENTS
This s udy was unded by Uni e si i Kebangsaan Malaysia and IRPA 09-02-
02-0001. We also wish o hank he managemen o Zoo Nega a and Uni e si i
PP- nninn Ulq17. n n nPi miooin-n n nn lii p.&P&n h nan l-i& n nnpi xT
LITERATURE CITED
BAKER, H. G. & HARRIS, B. J. 1957. The pollina ion o Pa kia by ba s and i s a endan e olu iona y
p oblems. E olu ion 11:449-460.
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