Vege a ion de e mines bu e ly di e si y and composi ion ac oss he
A abuko-Sokoke coas al o es in Kenya, a opical biodi e si y ho spo
Ma ia Fungomeli1, Ma in Wieme s2, Lucia Calde ini3, Alessand o Chia ucci3
1 Coas alFo es sConse a ionUni ,Cen e o Biodi e si y,Na ionalMuseumso Kenya,P.OBox596Kili i,Kenya
2 Senckenbe g Ge man En omological Resea ch Ins i u e, Ebe swalde S . 90, 15374 Münchebe g, Ge many
3 Biogeog aphy and Mac oecology G oup, Depa men o Biological, Geological and En i onmen al Sciences, Alma Ma e S udio um – Uni e si y
o Bologna, ia I ne io 42, 40126 Bologna, I aly
h ps://zoobank.o g/7CF5CEB5-FD82-49CC-BD5A-3574223248E9
Co esponding au ho : Ma ia Fungomeli ([email p o ec ed])
Academic edi o : Thomas Schmi | Recei ed 6 Ap il 2025 | Accep ed 31 Oc obe 2025 | Published 25 No embe 2025
Abs ac
Communi y s uc u es, including bu e ly di e si y, a e shaped by bo h bio ic and abio ic ac o s, wi h o es ype exe ing a
signi ican in luence. The A abuko Sokoke Fo es (ASF), he la ges emaining coas al o es agmen in Kenya and Eas A ica,
is ich in biodi e si y and endemic species. Gi en i s a ied o es ypes, ASF p o ides a unique oppo uni y o examine how hese
di e ences a ec bu e ly communi y s uc u e. This s udy aims o in es iga e how ege a ion di e si y and s uc u e in luence
bu e ly communi y s uc u es and species ichness wi hin ASF. We conduc ed bu e ly and woody plan su eys du ing he d y
season ac oss ou dis inc o es ypes in ASF: Cynome a o es , B achys egia woodland, mixed o es and he o es edge. Bu e ly
popula ions we e sampled using ansec s measu ing 10 m × 100 m and woody plan species we e su eyed along o e lapping
ansec s. A o al o 6,050 bu e ly indi iduals we e eco ded, ep esen ing 86 species ac oss 38 gene a and i e amilies. The
woody ege a ion comp ised 178 species, belonging o 78 gene a and 34 amilies. Signi ican di e ences in bu e ly species
abundance we e obse ed ac oss he o es ypes, hough no signi ican di e ences we e ound in species ichness. Be a di e si y
analyses e ealed consis en ly high communi y dissimila i y ac oss all o es ypes, d i en p edominan ly by balanced a ia ion in
species abundances a he han nes edness. B achys egia o es exhibi ed he highes o al be a di e si y, while o es edge exhibi ed
he lowes . This indica es ha species u no e , a he han ichness di e ences, is he p ima y mechanism s uc u ing bu e ly
communi ies a he landscape scale in A abuko Sokoke Fo es . Bu e ly species di e si y showed a s ong co ela ion wi h plan
species di e si y. Addi ionally, bu e ly wingspan size a ied signi ican ly amongs o es ypes. Ou indings unde sco e he c ucial
ole o na u al plan o es di e si y in suppo ing bu e ly di e si y and highligh he syne gis ic unc ions o he mixed o es and
B achys egia o es as key habi a s. The e is need o conse a ion s a egies ha accoun o mul iple dimensions o biodi e si y.
While mixed o es se es as a ese oi o high species ichness and abundance, B achys egia o es o e s c i ical alue h ough
hei con ibu ion o be a di e si y a he landscape le el. These esul s highligh he undamen al impo ance o conse a ion e o s
di ec ed o p o ec high plan di e si y and s uc u al he e ogenei y o p o ide a b oad spec um o ecological niches and habi a
connec i i y o bu e lies. Such s a egies will enhance bu e ly di e si y and con ibu e o e ec i e conse a ion in agmen ed
o es s and especially in A abuko Sokoke Fo es .
Key Wo ds
A abuko Sokoke Fo es , bu e ly di e si y, communi y s uc u e, o es edge, habi a connec i i y, habi a quali y, plan species,
species co-occu ence, opical o es s
Con ibu ions o En omology 75 (2) 2025, 299–318|DOI 10.3897/con ib.en omol.75.e155016
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use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
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Ma ia Fungomeli e al.: Bu e ly and ege a ion di e si y in e ac ions in A abuko Sokoke Fo es Kenya300
In oduc ion
Communi y s uc u es a e shaped by he in e play o
bio ic and abio ic ac o s, wi h o es ype exe ing a
signi ican in luence on bu e ly di e si y. Di e en
o es ypes can signi ican ly impac bu e ly di e si y
h ough changes in mic oclima e, esou ce a ailabili y,
plan di e si y and ege a ion s uc u e (Schwei ze and
Dey 2011). Bu e lies a e widely ega ded as e ec i e
bioindica o s o habi a quali y and ecosys em heal h due
o hei sensi i i y o en i onmen al changes and close
associa ions wi h hos plan s and mic ohabi a s (Bouye
e al. 2007; Dobson 2012). Thei communi y composi ion
and species ichness o en e lec unde lying pa e ns in
ege a ion di e si y and s uc u e, making hem aluable
o moni o ing biodi e si y esponses o habi a a ia ion.
Globally, an es ima ed 18,000 bu e ly species ha e
been documen ed, wi h app oxima ely 3,600 species
occu ing in A ica and a ound 870 eco ded in Kenya
(La sen 1991). The A abuko-Sokoke Fo es alone suppo s
o e 300 o Kenya’s bu e ly species, highligh ing i s
signi icance as a global biodi e si y ho spo (Ayiemba
1995; ASF Managemen Team 2002). I is es ima ed ha ,
globally, app oxima ely 90% o bu e lies a e ound in
opical a eas, bu hei ecological ole is less s udied han in
empe a e egions, which also applies o ege a ion s udies
(Boneb ake e al. 2010; see Fungomeli e al. (2020b)).
Bu e lies play a c ucial ole as a biogeog aphical
and ecological indica o g oup o habi a agmen a ion,
an h opogenic dis u bance and clima e change e ec s (La sen
1993; Heikkinen e al. 2009; Manzoo e al. 2013). Thei
li e cycle highly depends on plan s ei he o b eeding (hos
plan s) o ood (nec a ) and mul iple o he en i onmen al
ac o s (Collinge e al. 2003; Manzoo e al. 2013). They
can se e as indica o s o biodi e si y in ecological s udies
due o hei sensi i i y o e en mino changes in habi a
condi ions o dis u bances (Lomo e al. 2006; Bouye e al.
2007; Dobson 2012). In addi ion, bu e lies play an essen ial
ecological ole as pollina o s and he bi o es (Cou ney e al.
1982; Boneb ake e al. 2010; Rade e al. 2015). He bi o y
has been alued as a mechanism ha has p omo ed plan
co-exis ence and di e si y, while pollina ion has enhanced
plan s li e, g ow h and di e si y (Vail 1992; Coley and
Ba one 1996; Viola e al. 2010). Mo eo e , hei associa ion
wi h pa icula o es ypes and hos plan s, he luc ua ion
in hei ichness and abundance acco ding o seasonali y
and hei pe asi e p esence on he e i o y makes hem
pe ec s udy subjec s o in es iga ing and moni o ing he
conse a ion s a us o ecosys ems (Lien 2007; Monas y skii
2007; Habel e al. 2018). This co-exis ence and in e play
be ween bu e lies and plan s o e s a unique undamen al
con ibu ion o ecosys em unc ioning while p esen ing a
huge po en ial in opical o es s biodi e si y moni o ing
(Humpden and Na han 2010).
Vege a ion di e si y enhances ecological complexi y
by inc easing he a ailabili y o nec a sou ces, la al hos
plan s and mic oclima ic niches (Vu e al. 2015). S uc u al
cha ac e is ics, such as canopy heigh , oliage densi y
and e ical s a i ica ion, u he in luence mic ohabi a s
and esou ce accessibili y, di ec ly impac ing bu e ly
o aging beha iou , o iposi ion and su i al (Collinge
e al. 2003). Consequen ly, a eas wi h high ege a ion
he e ogenei y a e o en associa ed wi h g ea e bu e ly
species ichness and mo e s able communi y s uc u es.
Despi e his well-es ablished ela ionship, he ex en o
which ege a ion di e si y and s uc u e shape bu e ly
assemblages emains poo ly documen ed in many opical
and sub opical ecosys ems. This is mo e p onounced
especially in Eas A ica, a egion known o i s ecological
he e ogenei y and high biodi e si y. The A abuko-Sokoke
Fo es (ASF) in coas al Kenya p esen s a unique oppo uni y
o in es iga e hese ela ionships, as i encompasses a
mosaic o dis inc o es ypes wi hin a ela i ely compac
landscape, enabling de ailed compa isons o bu e ly
communi y assembly ac oss en i onmen al g adien s. The
o es hos s ou bu e ly species endemic o he o es s
o Kenya and Tanzania: Ac aea ma uapa, Baliochila
la ima gina a, Baliochila s ygia and Cha axes blanda,
50 na ionally and globally a e plan species, h ee a e
endemic mammals and is home o 230 bi d species, 15
o which a e a e and endemic o he Kenyan coas (ASF
Managemen Team 2002). The o es also plays a c ucial
ole as a global eco- ou ism si e, while locally suppo ing
su i al o he o es adjacen indigenous people
li elihoods who depend on he o es o bu e ly a ming,
collec ing medicinal plan s and wood p oduc ion.
Mo eo e , al hough ASF is ich in plan di e si y and
bu e ly di e si y, li le is known abou hei in e ac ion.
Limi ed bu e ly s udies ca ied ou in ASF ha e looked
a he bu e ly di e si y ac oss he o es and o es ypes
o seasonali y in luence on bu e ly di e si y (Ayiemba
1995; Habel e al. 2018). Howe e , o ou knowledge,
he e is no s udy ha has ho oughly in es iga ed he
in luence o he plan species di e si y on bu e ly
di e si y in ASF. Mo eo e , he need o egula ly assess
and moni o i s con inued agmen a ion and biodi e si y
is he e o e undamen al o long- e m conse a ion
e o s (Aze ia e al. 2007; MacFa lane e al. 2015; Habel
e al. 2017; Busck-Lumhol and T eue 2018). In his
s udy, we in es iga e he ela ionships be ween ege a ion
s uc u e and bu e ly communi y s uc u e and species
di e si y wi hin he A abuko-Sokoke Fo es (ASF). This
was achie ed h ough sys ema ic sampling along 100-m
ansec s ac oss di e en o es ypes, conduc ed o e
a ou -mon h pe iod, inco po a ing a ange o bu e ly
unc ional ai s o enhance ecological in e p e a ion.
In pa icula , we in es iga e: (i) how he dominan
o es ypes in luence bu e ly species di e si y, compo-
si ion and abundance in ASF; (ii) how plan species di-
e si y in luence o co ela e wi h bu e ly di e si y and
composi ion and (iii) how bu e ly wingspan ai s a y
ac oss di e en o es ypes. We syn hesise hese esul s
o be e guide he conse a ion policy o mula ions o
sus ainable o es use and managemen o he o es , es-
pecially in he d y season when his s udy was conduc ed.
Con ibu ions o En omology 75 (2) 2025, 299–318 301
Ma e ials and me hods
S udy a ea and o es ypes
The A abuko Sokoke Fo es (ASF) is he la ges o es
agmen emaining wi hin he Kenyan coas al o es s co -
e ing an a ea o 42,000 ha, he second being Shimba Hills
Fo es (25,300 ha; Fig. 1A; Bu gess and Cla ke (2000);
Fungomeli e al. (2020a)). I is globally alued as a wo ld
biodi e si y ho spo o he Eas e n A c and Coas al Fo es s
o Kenya and Tanzania (Mye s e al. 2000). ASF is a cen e
o endemism, hos ing a conspicuous numbe o h ea ened
and endange ed species and ecen ly decla ed a UNESCO
Biosphe e Rese e (UNESCO 2019). An h opogenic p es-
su e and biodi e si y loss, oge he wi h clima e change
a e hea ily impac ing opical o es s, such as he A abuko
Sokoke Fo es in Kenya (Bu gess and Cla ke 2000; New on
and Eche e ía 2014; FAO 2018; Fungomeli e al. 2025).
As a d y lowland coas al o es , ASF sp eads be ween
he ci ies o Kili i in he sou h and Malindi in he no h,
posi ioned be ween 39°48'E and 40°00'E longi ude and
be ween 3°11'S and 3°29'S la i ude (Fanshawe 1995;
Muchi i e al. 2001). I lies on a la coas al plain a sea
le el and he a ea is di ided by a low esca pmen which
c osses he o es om sou h-wes o no h-eas (Moomaw
1960; Fanshawe 1995).
The clima e consis s o ainy and d y seasons, wi h
wo ain all seasons o long and sho ains. The long
ainy season occu s om Ap il o July; sho ains om
Oc obe o Decembe , while he d y season las s om
Decembe o Ma ch and in Augus /Sep embe (Bu gess
and Cla ke 2000; Omenge 2002). The annual ain all
anges om 600 o 1,000 mm, wi h ain all dec easing
om eas o wes wi hin he Fo es (Omenge 2002; Habel
e al. 2017). Tempe a u e anges om mon hly a e ages
o 24 °C and 30 °C and humidi y is abou 60% annual-
ly (Bu gess and Cla ke 2000). Se e al wa e pools exis
wi hin he Fo es du ing he ainy season wi h mos d y-
ing ou in he d y season and he e a e no i e s wi hin
he o es (Fungomeli e al. 2001; Kanga 2002; Mu ii hi
and Kenyon 2002).
A de ining cha ac e is ic o he A abuko-Sokoke
Fo es is he p esence o dis inc ege a ion ypes,
usually e e ed o as o es ypes (Fig. 1B–E). These
o es ypes a e closely associa ed wi h he unde lying
soil cha ac e is ics. The a ea ea u es wo p edominan
soil ypes: ligh , whi e sandy soils and hea ie , ed clay
soils (Fanshawe 1995; Muchi i e al. 2001). These con-
as ing soil condi ions ha e signi ican ly in luenced
he dis ibu ion and composi ion o he o es ’s ege a-
ion communi ies. The ou dis inc o es ypes a e he
Cynome a o es , B achys egia o es , Mixed o es
and he o es edge (Fig. 1B–E).
Cynome a o es
The Cynome a o es , which occupies he wes e n
sec o o he ASF, is he mos ex ensi e o he o es
ypes, accoun ing o o e 50% o he o es a ea
(Fanshawe 1995). This zone is ound p edominan ly
on ed clay soils and is cha ac e ied by a dense, low-
s a u ed canopy, composed mainly o Cynome a
Figu e 1. A. Map o A abuko Sokoke Fo es , Kenya, showing he dis ibu ion o he 108 s udied bu e ly ansec s wi hin he ou o -
es ypes: B achys egia, Cynome a, Mixed o es and Fo es edge. B–E. The ou o es ypes s udy si es wi hin A abuko Sokoke Fo -
es , Kenya showing. B. Cynome a o es ; C. B achys egia o es ; D. Mixed o es ; E. Fo es edge. Pho o c edi s: Ma ia Fungomeli.
AB C
D E
Ma ia Fungomeli e al.: Bu e ly and ege a ion di e si y in e ac ions in A abuko Sokoke Fo es Kenya302
webbe i, Cynome a suhalensis and Manilka a
sulca a, wi h occasional eme gen s, such as Eupho bia
candelab um (Fig. 1). The unde g ow h is spa se due o
he closed canopy and low ligh pene a ion, c ea ing a
ela i ely s able mic oclima e wi h educed empe a u e
luc ua ions and high humidi y.
B achys egia o es
B achys egia o es is loca ed cen ally wi hin he ASF,
on he nu ien -poo , whi e sandy soils and co e s app ox-
ima ely 18% o he o es (Fig. 1; Muchi i e al. 2001). I
is domina ed by B achys egia spici o mis, cha ac e ised
by an open canopy s uc u e in e spe sed wi h g asses
and sh ubs, esul ing in a mo e sunli and d ie en i on-
men han o he o es ypes. The lo is ic composi ion is
adap ed o d y condi ions and he habi a is impo an o
bu e ly species, ep iles and bi ds ha depend on high
ligh a ailabili y and open unde s o eys. Addi ionally, he
openness o his zone c ea es mic ohabi a s a ou able
o he mo egula ion and basking, c ucial o many in-
e eb a e and he pe o auna species.
Mixed o es
The mixed o es ype is loca ed in he eas e n pa o
he Fo es , whe e i g ows on g ey sandy soils ha e-
ain mo e mois u e han he whi e sands, bu a e ligh e
han he ed clay soils. This o es ype ep esen s abou
17% o he Rese e and is no able o i s high plan spe-
cies ichness and e ical s a i ica ion (Fanshawe 1995;
Muchi i e al. 2001; A abuko Sokoke Managemen Team
2002). I is domina ed by mixed plan species o A zelia
quanzensis, Hymenaea e ucosa, New onia hildeb and-
ii and Manilka a sansiba ensis.
Fo es edge
The o es edge was selec ed along he ansi ion om he
mixed o es o he A abuko-Sokoke Fo es (ASF) o ad-
jacen ag icul u al lands. This eco one ep esen s a g ad-
ual shi om in ac o es ecosys ems o human-modi-
ied landscapes and coas al habi a s (Fig. 1; Muchi i e al.
2001). I is cha ac e ised by a he e ogeneous mosaic o
land uses, including subsis ence a ms, sca e ed sh ubs
and open g assy clea ings. S uc u ally, i exhibi s in-
c eased ligh pene a ion, highe empe a u e luc ua ions
and educed canopy co e compa ed o he o es in e io .
These condi ions c ea e a dis inc i e mic oclima e ha
suppo s a unique assemblage o lo a and auna adap ed
o edge en i onmen s.
Da a collec ion
Field sampling and da a collec ion we e conduc ed
du ing he d y season mon hs (Janua y-Ap il) o 2019
ac oss he ou o es ypes o ASF: Cynome a o es ,
B achys egia woodland, mixed o es and o es edge
(Fig. 1). Bu e lies we e sampled by using a s anda d
numbe o 27 ansec s wi hin each ege a ion ype
leading o a o al o 108 ansec s. Each ansec mea-
su ed 10 m × 100 m. Bu e lies we e eco ded in each
ansec by using a s anda d coun echnique pe o med
by walking a slow cons an pace o app oxima ely
15 min. All bu e ly species seen on bo h sides o he
pa h we e eco ded. Each ansec was su eyed once
pe day, e e y day, h oughou he ou -mon h d y sea-
son (Janua y-Ap il 2019), ensu ing comp ehensi e and
exhaus i e species de ec ion in acco dance wi h es ab-
lished bu e ly moni o ing p o ocols (Polla d 1977;
Polla d and Ya es 1993).
Bu e lies we e iden i ied and eco ded a species le -
el. Specimens ha could no immedia ely be iden i ied
in he ield we e caugh wi h a bu e ly ne and placed
in numbe ed en elopes o pho og aphed o u he iden-
i ica ion in he lab. Iden i ica ion was ca ied ou using
he bu e ly e e ences o he a ea (La sen 1991) and
suppo ed by axonomic coun e checks om published
sou ces. All ansec s we e geo- e e enced, wi h de ails
o da e, hou o s a and end.
Vege a ion ield sampling was pe o med by using 27
plo s each measu ing 10 m × 100 m (same used o bu e -
ly ansec s he ea e e e ed o as plo s) and in e nally
subdi ided in o 20 subplo s o 10 m × 5 m. Each eg-
e a ion plo co esponded o a bu e ly ansec . Wi hin
he plo s and subplo s, we iden i ied and measu ed he
heigh and diame e a b eas heigh (DBH) o each indi-
idual woody plan species ( ees, lianas and sh ub) wi h
DBH ≥ 5 cm. Plan s wi h DBH < 5 cm, such as small
sh ubs, we e iden i ied in wo subplo s o each plo (see
Fungomeli e al. 2020a, 2020b).
Bu e ly ai s: wingspan sizes
We compiled and ob ained wingspan sizes o ou sampled
bu e ly species om published da a sou ces o Woodhall
(2005), Woodhall (2020), Schmi (pe s. comm), Ba code
o Li e Da a Sys em da abase (h ps:// 3.boldsys ems.
o g) and om he collec ion o he Senckenbe g Ge man
En omological Ins i u e, Münchebe g.
All bu e lies encoun e ed along he ansec s we e
classi ied acco ding o hei ecological ai s and dis i-
bu ion. The la al die o each species was de e mined,
based on hos plan use and ca ego ised in o one o h ee
ophic b ead h classes: (1) monophagous, es ic ed o
a single hos plan genus; (2) oligophagous, es ic ed o
hos plan s wi hin a single plan amily; o (3) polypha-
gous, u ilising hos plan s om mul iple plan amilies. A
u he classi ica ion in o endemic s a us was assigned o
species acco ding o La sen (1996).
Da a analysis
A communi y ma ix was p epa ed o he bu e ly spe-
cies abundances and ano he ma ix was p epa ed o he
woody plan species ac oss he o es ypes.
Con ibu ions o En omology 75 (2) 2025, 299–318 303
Bu e ly species di e si y
Bu e ly species di e si y was analysed in e ms o spe-
cies ichness, Shannon index and Simpson index ac oss
o es ypes:
Shannon Index:
Simpson Index:
Fo bo h indices, k ep esen s he o al numbe o spe-
cies, while pi indica es he ela i e abundance o each
species ha is calcula ed as ni/N (in which ni indica es he
numbe o indi iduals o he i-species and N indica es he
numbe o indi iduals o all species wi hin he ansec .
Bu e ly species ichness and mean abundance dis-
ibu ions ac oss o es ypes we e isualised using box-
plo s. Mann-Whi ney U es o pai wise compa isons was
used o compa e species ichness and abundance amongs
o es ypes. Addi ionally, ank-abundance cu es we e
cons uc ed o each o es ype o illus a e pa e ns o
species dominance and e enness (Whi ake 1965).
Ra e ac ion cu es and species di e si y es ima ion
To assess and compa e species di e si y ac oss o es
ypes, we employed sample co e age-based a e ac ion and
ex apola ion wi hin he Hill numbe s amewo k (Chao e
al. 2014). We analysed abundance da a o each o es ype
using he iNEXT() unc ion om he iNEXT package in
R (Hsieh e al. 2016), speci ying da a ype = “abundance”.
This unc ion pe o ms bo h in e pola ion ( a e ac ion) and
ex apola ion o di e si y es ima es, based on indi idual-
based abundance da a. Con idence in e als (95%) we e
de i ed using 1,000 boo s ap eplica es. Following Chao e
al. (2014), ex apola ion was cons ained o a maximum o
wice he e e ence sample size o main ain es ima e s abili y.
Co ela ion be ween bu e ly and plan species di e si y
We applied a symme ic Co-co espondence analysis
(CoCA) o quan i y ela ionships be ween he plan species
communi y wi h he bu e ly species communi y ac oss he
o es ypes. Co-co espondence analysis is use ul o com-
pa ing biological communi ies whe e obse a ions ha e
been made a he same loca ions (B aak and Scha e s 2004).
We did his by a weigh ed a e age o species abundance
alues o plan species and sepa a ely o bu e ly species
wi hin each o he o es ypes. We used ‘coca’ unc ion
o he ‘coco esp’ R package (Simpson 2009) o co ela e
he bu e ly and plan communi ies using he ‘symme ic’
me hod. All g aph plo ing was pe o med using R package
ggplo 2 (Wickham 2016) and gg epel (Slowikowski 2020).
Bu e ly species composi ion
We squa e- oo ans o med bu e ly communi y abun-
dances p io o he analysis o educe e ec s o dominan
species. T ans o med communi y abundances we e hen
used o gene a e a B ay-Cu is dissimila i y ma ix (B ay
and Cu is 1957). We es ed o species composi ion di e -
ences in he bu e ly communi y s uc u e amongs o es
ypes by an analysis o simila i ies (ANOSIM) using he
‘anosim’ unc ion o he ‘ egan’ R package (Oksanen e al.
2020). We also es ed o signi ican di e ences be ween
o es ypes using he pe mu a ional analysis o a iance
(PERMANOVA), using he ‘adonis’ unc ion o he ‘ eg-
an’ R package. All es s we e conduc ed using 999 pe mu-
a ions. Bu e ly species con ibu ing o simila i ies ac oss
o es ypes we e de e mined using simila i y pe cen ag-
es analysis (SIMPER). P- alues we e adjus ed using he
Benjamini-Hochbe g es o con ol he alse disco e y
a e. This app oach was selec ed o i s abili y o limi ype
I e o s, while e aining g ea e s a is ical powe han mo e
conse a i e me hods, such as he Bon e oni co ec ion.
Be a di e si y pa i ioning
To assess whe he a iabili y in bu e ly communi y
composi ion di e ed amongs he ou o es ypes
(B achys egia, Cynome a, mixed o es and o es edge),
we pe o med a pe mu a ion es o homogenei y o
mul i a ia e dispe sions (PERMDISP), based on B ay-
Cu is dissimila i ies using he ‘be adispe ’ unc ion in
he egan R package. We hen e alua ed be a di e si y
amongs he ou o es ypes. We quan i ied mul i-si e
be a di e si y using he abundance-based ex ension
o he ‘be apa ’ amewo k (Baselga e al. 2017),
implemen ed ia he ‘be a.mul i.abund’ unc ion in he
R package be apa ( . 1.6). This me hod pa i ions B ay-
Cu is dissimila i y in o h ee componen s: (i) o al be a
di e si y (β_ o al); (ii) balanced a ia ion in abundance
(β_balanced) and (iii) abundance g adien s (β_g adien ).
We also assessed spa ial he e ogenei y o communi y
composi ion wi hin o es ypes by es ing homogenei y
o mul i a ia e dispe sion. B ay-Cu is dissimila i ies,
calcula ed om species abundance da a, we e used o
compu e dis ances o indi idual plo s o hei espec i e
g oup cen oids using he ‘be adispe ’ unc ion in he R
package egan (Oksanen e al. 2020). These dis ances e-
lec wi hin-g oup a ia ion in communi y composi ion.
Di e ences in dispe sion amongs o es ypes we e es -
ed using ANOVA, ollowed by Tukey’s Hones Signi i-
can Di e ence (HSD) pos -hoc es s.
Bu e ly composi ion ‒ NMDS
To isualise di e ences in bu e ly species composi-
ion amongs o es ypes, we conduc ed a non-me ic
mul idimensional scaling (NMDS) analysis, based on
B ay-Cu is dissimila i ies (K uskal 1964). P io o anal-
ysis, bu e ly species abundance da a we e squa e- oo
ans o med o educe he in luence o highly abundan
species. NMDS was pe o med using he ‘me aMDS’
unc ion om he egan package (Oksanen e al. 2020) in
R (R Co e Team 2020). This unc ion conduc s au oma ic
da a s anda disa ion, mul iple andom s a s and i e a ion
Ma ia Fungomeli e al.: Bu e ly and ege a ion di e si y in e ac ions in A abuko Sokoke Fo es Kenya304
o ensu e a s able and op imal o dina ion solu ion. G oup-
ings by o es ype we e isualised using con ex hulls.
Bu e ly ai s: wingspan sizes
Using Pea son co ela ion, we co ela ed bu e ly wing-
span sizes ac oss o es ypes, by i s co ela ing o o al
abundances in all o es ypes and hen second wi hin each
ege a ion ype. Following con i ma ion o no mali y,
pai wise - es s we e conduc ed o compa e a e age wing-
span sizes ac oss di e en o es ypes. To accoun o
mul iple compa isons and con ol he alse disco e y a e,
p- alues we e adjus ed using he Benjamin-Hochbe g es .
Resul s
We eco ded a o al o 6,050 bu e ly indi iduals belong-
ing o 86 species, 38 gene a and i e amilies ac oss he
ou o es ypes o A abuko Sokoke Fo es (Appendix 1).
The plan species su ey esul ed in a o al o 178 plan
species belonging o 78 gene a and 34 amilies.
Bu e ly species di e si y
Bu e ly species di e si y was p ima ily domina ed by
he Nymphalidae amily, which had he highes numbe
o species, ollowed by Pie idae, Papilionidae, Lycaeni-
dae and Hespe iidae (Appendix 1). Analysis on he mos
abundan bu e ly species e ealed Phalan a phalan ha,
Appias epaphia, Ca opsilia lo ella, Hypolimnas misip-
pus and Coeliades o es an, as he mos equen ac oss
all o es ypes (Suppl. ma e ial 1: ig. S1). A s ong pos-
i i e co ela ion was obse ed be ween species ichness
and abundance ac oss he o es ypes (R² = 0.89). The
dis ibu ion o bu e ly la al eeding habi s showed oli-
gophagous and polyphagous species being dominan in
all o es ypes (Suppl. ma e ial 1: ig. S2).
Ra e ac ion cu es and species di e si y es ima ion
Ra e ac ion cu es e ealed he mixed o es exhibi ed
he highes species ichness, ollowed by B achys egia
o es , o es edge and Cynome a o es s (Fig. 2).
Ra e ac ion cu es o all o es ypes app oached
asymp o es, indica ing su icien sampling and species
ichness cap u e, excep in he B achys egia o es , whe e
he non-asymp o ic cu e sugges s ha u he sampling
may unco e addi ional species.
Bu e ly species ichness and abundances ac oss o es
ypes showed ha he mixed o es had he highes cumu-
la i e species ichness, ollowed by B achys egia and o -
es edge, while Cynome a had he lowes alue (Table 1).
A e age species ichness and abundance pe plo a ied
ac oss o es ypes, wi h signi ican ly highe abundance
alues obse ed a he o es edge and in mixed o es hab-
i a s (ANOVA, P < 0.05; Fig. 3). Acco ding o ANOVA,
di e ences amongs species ichness pe ansec we e a
he signi icance h eshold amongs o es ypes (P = 0.05),
while species abundances pe ansec we e signi ican ly
di e en (P = 0.001; Fig. 3). Pai wise compa isons o bu -
e ly abundance e ealed signi ican di e ences be ween
Cynome a o es and o es edge (P = 0.001), B achys e-
gia o es and o es edge (P = 0.013), as well as be ween
B achys egia o es and o es edge (P = 0.001; Fig. 3).
The di e si y indices indica ed ela i ely simila le els
o bu e ly di e si y ac oss ege a ion ypes. Shannon in-
dex alues anged om 2.91 ± 0.40 a he o es edge o
2.82 ± 0.42 in he mixed o es , while he Simpson index
alues anged om 0.93 ± 0.04 o 0.92 ± 0.03.
Be a di e si y pa i ioning
Mul i a ia e dispe sions e ealed signi ican di e -
ence in mul i a ia e dispe sion ac oss he ou o es
ypes (F = 3.893, P = 0.007). The e o e, a ia ion in
bu e ly communi y composi ion may in pa be in-
luenced by di e ences in wi hin- o es ype he e o-
genei y. Mul i- o es ype be a di e si y analysis (β)
e ealed consis en ly high o al dissimila i y ac oss
o es ypes, wi h β_ o al alues anging om 0.885 o
Table 1. The bu e ly species di e si y ac oss o es ypes,
showing cumula i e species ichness and abundance, Shan-
non index and Simpson index pe ege a ion ype in A abuko
Sokoke Fo es , Kenya.
Species di e si y B achys egia Cynome a Fo es
edge
Mixed
o es
Cumula i e species
ichness
50 40 52 80
Cumula i e species
abundance
1022 1112 2141 1775
Shannon’s H Index 2.38 2.58 2.87 2.66
Simpson’s 1-D Index 0.9 0.9 0.93 0.91
Figu e 2. Ra e ac ion cu es showing species ichness as a
unc ion o numbe o indi iduals ac oss he sampled o es
ypes o B achys egia, Cynome a, Mixed o es and Fo es
edge in he A abuko Sokoke o es , Kenya. Each solid line ep-
esen s ac ual sampled species (in e pola ed), and he dashed-
line ep esen s ex apola ed indi iduals (ex apola ed). Shaded
a eas ep esen 95% con idence in e al.
Con ibu ions o En omology 75 (2) 2025, 299–318 305
0.910 (Table 2). The la ges con ibu ion o be a di e -
si y came om balanced a ia ion in species abundance
(β_balanced: 0.803‒0.837), indica ing ha communi y
composi ional di e ences we e p ima ily d i en by
species u no e in abundance a he han nes edness
(β_g adien : 0.067‒0.096) (Table 2). B achys egia o -
es exhibi ed he highes o al be a di e si y (β_ o al =
0.910), sugges ing subs an ial he e ogenei y in commu-
ni y composi ion ac oss plo s. Fo es edge had he low-
es β_ o al (0.885), while Cynome a and Mixed o es s
showed in e media e alues (Table 2).
Spa ial dispe sion o communi y composi ion
Spa ial he e ogenei y wi hin o es ypes, assessed ia
mul i a ia e dispe sion, a ied signi ican ly (Table 2).
B achys egia o es exhibi ed he highes dispe sion
(mean dis ance = 0.45), indica ing g ea e spa ial a i-
abili y in species composi ion. These esul s e lec sub-
s an ial spa ial he e ogenei y in species abundances in
B achys egia. In con as , o es edge had he lowes dis-
pe sion (median ~ 0.33), indica ing mo e homogeneous
communi y s uc u e. Tukey HSD es s con i med dispe -
sion in B achys egia was signi ican ly g ea e han a o -
es edges (P = 0.006), while o he pai wise compa isons
we e no s a is ically signi ican (Table 2). These esul s
e lec ha o es edge communi ies a e mo e simila o
each o he , while B achys egia o es is mo e ecologically
di e se. O e all, be a di e si y esul s indica e ha abun-
dance-based species u no e d i es communi y di e en-
ia ion ac oss o es ypes, wi h B achys egia o es s sup-
po ing mo e spa ially he e ogeneous communi ies, while
o es edges ha bou mo e homogenised assemblages.
Co ela ion be ween bu e ly and plan species
di e si y
Co-co espondence analysis (CoCA) e ealed a s ong
co ela ion be ween he species composi ion o plan s and
bu e lies ac oss he o es ypes. The co ela ion coe -
icien s o Axis 1 and Axis 2 be ween he bu e ly and
plan communi ies we e 0.991 and 0.994, espec i ely.
The eigen alues o he i s and second axes indica ed
he con ibu ion o each axis o he o al ine ia, wi h al-
ues o 0.022 and 0.012, ep esen ing a a iance o 57.3%
and 32.6%, espec i ely. This esul ed in a o al explained
a iance o 89.9% (Fig. 4), highligh ing a obus and
highly signi ican co ela ion be ween he communi y
ma ices o plan s and bu e lies.
Bu e ly species composi ion
The NMDS analysis o bu e ly species composi ion
ac oss o es ypes e ealed conside able o e lap, wi h
no clea sepa a ion obse ed amongs he di e en o -
es ypes, wi h Cynome a o es co e ing a wide NMDS
space ha o e laps B achys egia, o es edge and mixed
o es (Fig. 5, Table 2). Bu e ly assemblages ac oss di -
e en o es ypes showed subs an ial o e lap, wi h many
species occu ing in mo e han one o es ype wi hin
he ASF. A pai wise pe mu a ional mul i a ia e analysis
o a iance (PERMANOVA) e ealed s a is ically sig-
ni ican composi ional di e ences in bu e ly assem-
Figu e 3. Boxplo s showing bu e ly species ichness and
abundances compa ison ac oss he ou o es ypes o B achys-
egia, Cynome a, Fo es edge and Mixed o es in he A abuko
Sokoke Fo es , Kenya. Fo es ype wi h di e en le e deno es
s a is ical signi ican di e ences (P < 0.01).
Table 2. Summa y o be a di e si y me ics ac oss he ou o es ypes, including abundance-based pa i ioning componen s o o al
be a di e si y (β_ o al); balanced a ia ion (β_balanced); and abundance g adien (β_g adien ), as well as mul i a ia e dispe sion
(measu ed by he mean dis ance o cen oid. Tukey HSD pos hoc es s we e used o pai wise compa isons amongs o es ypes
(n.s. = no signi ican di e ence).
Fo es ype n(plo s) β_ o al β_balanced β_g adien Mean dispe sion (dis ance o cen oid) Signi ican di e ence (Tukey HSD)
B achys egia 27 0.910 0.825 0.085 0.452 highe han o es edge (P = 0.006)
Cynome a 27 0.903 0.837 0.067 0.425 n.s. s. o he ypes
Mixed o es 27 0.903 0.807 0.096 0.418 n.s. s. o he ypes
Fo es edge 27 0.885 0.803 0.083 0.364 Lowe han B achys egia (P = 0.006)
Ma ia Fungomeli e al.: Bu e ly and ege a ion di e si y in e ac ions in A abuko Sokoke Fo es Kenya306
Figu e 4. Symme ic co-co espondence analysis (CoCA) o dina ion bi-plo s showing co ela ions be ween (a) bu e ly and plan
species and (b) plan and bu e ly species wi hin he ou o es ypes o B achys egia, Cynome a, Mixed o es and Fo es edge
in A abuko Sokoke Fo es , Kenya. The Axis-1 eigen alue o 0.022 explains a a iance o 57.3% and Axis-2 eigen alue o 0.012
explains a a iance o 32.6%. To al explained a iance by Axis 1 and 2 is 89.9%.
Figu e 5. Non-me ic mul idimensional scaling (NMDS) o
bu e ly species composi ion wi hin he ou o es ypes o
A abuko Sokoke Fo es , Kenya. Di e en colou s ep esen di -
e en o es ypes as ollows: B achys egia ( ed), Cynome a
(blue), Mixed o es (g een), Fo es edge (yellow).
Figu e 6. Bu e ly a e age wingspan sizes ac oss he ou o -
es ypes o B achys egia, Cynome a, Mixed o es and Fo es
edge in A abuko Sokoke Fo es . Fo es ype wi h di e en de-
no es signi ican di e ence (P < 0.01).
Con ibu ions o En omology 75 (2) 2025, 299–318 307
blages amongs he o es ypes (R2 = 0.07; P = 0.006).
Addi ionally, SIMPER esul s show species composi ion
di e ences amongs o es ypes ha con ibu e up o
70% o he obse ed dissimila i ies (Appendix 2).
Bu e ly ai s: wingspan sizes
A e age wingspan sizes we e signi ican ly la ge a he
o es edge compa ed o he Cynome a o es (P < 0.01;
Fig. 6). Howe e , no signi ican co ela ion was ound be-
ween wingspan size and species abundance ac oss o es
ypes (Fig. 7).
Bu e ly ai s: la al eeding habi s
The majo i y o la ae om he species encoun e ed we e
classi ied as oligophagous, ollowed by polyphagous
species and a smalle numbe classi ied as monophagous
(Suppl. ma e ial 1: ig. S2). O e all, Oligophagous spe-
cies cons i u ed he la ges sha e o indi iduals ac oss all
o es ypes, accoun ing o 63.4% o bu e lies a he
o es edge and eaching up o 67.5% in he mixed o es
(Suppl. ma e ial 1: ig. S2).
Discussion
This s udy in es iga ed he in luence o ege a ion di-
e si y and s uc u al complexi y on bu e ly commu-
ni y composi ion and species ichness wi hin A abuko
Sokoke Fo es (ASF), a coas al biodi e si y ho spo in
Eas A ica. Ou esul s p o ide new insigh s in o bu e -
ly communi y composi ion ac oss habi a ypes wi hin
he A abuko Sokoke Fo es and hei associa ions wi h
plan communi ies. By examining species dis ibu ions
alongside ege a ion da a, we highligh bo h b oad and
ine-scale pa e ns ele an o biodi e si y conse a ion
in opical o es mosaics.
Figu e 7. Bu e ly wingspan sizes co ela ion ac oss he ou o es ypes o A abuko Sokoke Fo es , Kenya. Showing wingspan
co ela ion o (a) The o al co ela ion in he ou o es ypes (b) he co ela ion o each o es ype o B achys egia, Cynome a,
Mixed o es and Fo es edge.
Ma ia Fungomeli e al.: Bu e ly and ege a ion di e si y in e ac ions in A abuko Sokoke Fo es Kenya314
Species Family
Cha axes jahlusa T imen, 1862 Nymphalidae
Cha axes jasius sa u nus Bu le , 1866 Nymphalidae
Cha axes las i G ose-Smi h, 1889 Nymphalidae
Cha axes p o oclea Feis hamel, 1850 Nymphalidae
Cha axes sp. Nymphalidae
Cha axes a anes (C ame , 1764) Nymphalidae
Cha axes iole a G ose-Smi h, 1885 Nymphalidae
Cha axes zoolina Wes wood, 1850 Nymphalidae
Coeliades o es an S oll, 1782 Hespe iidae
Colo is ama a (Fab icius, 1775) Pie idae
Colo is auxo (Lucas, 1852) Pie idae
Colo is danae (Fab icius, 1775) Pie idae
Colo is e is (Klug, 1829) Pie idae
Colo is euippe (Linnaeus, 1758) Pie idae
Colo is ione (Goda , 1819) Pie idae
Colo is p o omedia (Klug, 1829) Pie idae
Colo is egina (T imen, 1863) Pie idae
Colo is es a (Reiche, 1850) Pie idae
Cupidopsis ioba es (Hop e , 1855) Lycaenidae
Danaus ch ysippus do ippus Klug, 1845 Nymphalidae
Dixeia cha ina (Boisdu al, 1836) Pie idae
E onia cleodo a Hübne , 1823 Pie idae
Euphaed a neoph on Hop e , 1855 Nymphalidae
Eu ema sp. Pie idae
Eu y ela d yope C ame , 1779 Nymphalidae
Euxan hewake ieldi (Wa d, 1873) Nymphalidae
G aphium angolanus (Goeze, 1779) Papilionidae
G aphium an heus (C ame , 1779) Papilionidae
G aphium colonna (Wa d, 1873) Papilionidae
G aphium ki byi (Hewi son, 1872) Papilionidae
G aphium leonidas (Fab icius, 1793) Papilionidae
G aphium philonoe (Wa d, 1873) Papilionidae
G aphium policenes (C ame , 1775) Papilionidae
G aphium polis a us (G ose-Smi h, 1889) Papilionidae
G aphium po haon (Hewi son, 1865) Papilionidae
Ha ma heobene Doubleday, [1848] Nymphalidae
Hypolimnas an hedon (Doubleday, 1845) Nymphalidae
Hypolimnas decep o T imen, 1873 Nymphalidae
Hypolimnas misippus (Linnaeus, 1764) Nymphalidae
Junonia hie a (Fab icius, 1798) Nymphalidae
Junonia na alica Felde , 1860 Nymphalidae
Junonia oenone Linnaeus, 1764 Nymphalidae
Lep osia alces a (S oll, [1782]) Pie idae
Liby hea labdaca Wes wood, 1851 Nymphalidae
Melani is leda Linnaeus, 1758 Nymphalidae
Mylo h is aga hina (C ame , 1779) Pie idae
Nephe onia halassina (Boisdu al, 1836) Pie idae
Nep is sp. Nymphalidae
Papilio cons an inus Wa d, 1871 Papilionidae
Papilio da danus B own, 1776 Papilionidae
Papilio demodocus Espe , 1798 Papilionidae
Papilio ni eus Linnaeus, 1758 Papilionidae
Pa dopsis punc a issima Boisdu al, 1833 Nymphalidae
Phalan a phalan ha D u y, 1773 Nymphalidae
Physcaeneu a leda Ge s aecke , 1871 Nymphalidae
Pinacop e yx e iphia (Goda , 1819) Pie idae
Pseudac aea boisdu ali (Doubleday, 1845) Nymphalidae
Pseudac aea luc e ia (C ame , 1775) Nymphalidae
Salamis anaca dii Linnaeus, 1758 Nymphalidae
Salamis pa hassus D u y, 1782 Nymphalidae
Ti umala pe i e ana Doubleday, 1847 Nymphalidae
Con ibu ions o En omology 75 (2) 2025, 299–318 315
Appendix 2
SIMPER analysis o bu e ly species composi ion dissimila i ies esul s. Highligh ed a e species ha cumula i ely
con ibu e up o 70% o he obse ed dissimila i ies.
Table A2. Bu e ly species con ibu ing up o 70% o he obse ed dissimila i ies ac oss he sampled o es ypes o B achys egia,
Cynome a, Mixed o es and Fo es edge in he A abuko Sokoke o es , Kenya. Whe e a e age = he a e age con ibu ion o a
species o he dissimila i y be ween g oups; Sd = s anda d de ia ion o he species’ con ibu ion, showing a iabili y in how much
ha species con ibu es ac oss di e en sample compa isons; Ra io = he a e age con ibu ion di ided by i s s anda d de ia ion
(a e age / sd). A highe a io indica es ha he species consis en ly con ibu es o dissimila i y (less a iable); a a = he a e age
abundance o alue o he species in o es A (e.g. B achys egia); a b = he a e age abundance o alue o he species in o es ype
B (e.g. Cynome a); cumsum = he cumula i e sum o he con ibu ions up o he cu en species, usually exp essed as a pe cen age
o o al dissimila i y explained so a . This helps iden i y which species collec i ely con ibu e o a speci ied h eshold (e.g., 70%);
P- alue = s a is ical signi icance es ing he con ibu ion o ha species o he dissimila i y, lowe alues ( ypically < 0.05) sugges
he species con ibu es signi ican ly o di e ences be ween g oups.
Species a e age sd a io a a a b Cumsum % o o al dissimila i y P- alue
B achys egia s Cynome a
Phalan a phalan ha 0.030 0.022 1.330 2.143 1.564 0.049 4.9 0.001
Ca opsilia lo ella 0.027 0.023 1.160 1.234 1.646 0.094 9.4 0.002
Appias epaphia 0.026 0.022 1.147 1.549 1.828 0.137 13.7 0.001
Hypolimnas misippus 0.025 0.021 1.184 0.959 1.151 0.178 17.8 0.002
Colo is egina 0.023 0.021 1.121 0.397 1.122 0.217 21.7 0.001
G aphium philonoe 0.022 0.019 1.181 0.606 1.190 0.254 25.4 0.001
G aphium an heus 0.022 0.020 1.112 1.010 1.018 0.291 29.1 0.002
Papilio demodocus 0.022 0.018 1.202 1.106 0.977 0.327 32.7 0.006
Nep is sp. 0.022 0.019 1.154 0.884 1.001 0.363 36.3 0.001
Junonia oenone 0.020 0.017 1.195 0.958 0.841 0.397 39.7 0.031
Colo is euippe 0.018 0.018 1.034 0.784 0.569 0.427 42.7 0.025
E onia cleodo a 0.018 0.017 1.080 0.700 0.785 0.458 45.8 0.023
G aphium po haon 0.017 0.017 1.013 0.553 0.665 0.487 48.7 0.116
Colo is auxo 0.017 0.015 1.123 0.427 0.739 0.515 51.5 0.001
Hypolimnas decep o 0.017 0.018 0.968 0.000 0.741 0.544 54.4 0.001
Coeliades o es an 0.017 0.017 0.974 0.628 0.668 0.572 57.2 1.000
Ac aea sp. 0.017 0.016 1.072 0.612 0.729 0.600 60.0 0.006
Cha axes candiope 0.016 0.015 1.036 0.037 0.705 0.625 62.5 0.001
Pa dopsis punc a issima 0.015 0.016 0.946 0.552 0.501 0.651 65.1 0.001
Bicyclussa i za 0.014 0.016 0.910 0.154 0.616 0.675 67.5 0.001
Eu ema sp. 0.014 0.023 0.632 0.639 0.117 0.699 69.9 0.573
B achys egia s Fo es edge
Coeliades o es an 0.033 0.018 1.828 0.628 2.529 0.055 5.5 0.001
Hypolimnas misippus 0.021 0.015 1.364 0.959 1.738 0.090 9.0 0.183
Junonia oenone 0.021 0.015 1.331 0.958 1.743 0.124 12.4 0.022
Papilio demodocus 0.020 0.017 1.142 1.106 1.910 0.158 15.8 0.084
Ca opsilia lo ella 0.020 0.019 1.034 1.234 1.415 0.191 19.1 0.809
Phalan a phalan ha 0.019 0.023 0.846 2.143 2.479 0.224 22.4 0.941
Appias epaphia 0.019 0.018 1.063 1.549 2.052 0.255 25.5 0.489
Eu ema sp. 0.019 0.018 1.052 0.639 1.176 0.287 28.7 0.006
G aphium an heus 0.018 0.016 1.126 1.010 1.316 0.317 31.7 0.445
Colo is ione 0.018 0.016 1.126 0.295 1.185 0.347 34.7 0.001
Colo is euippe 0.017 0.015 1.144 0.784 1.144 0.375 37.5 0.261
E onia cleodo a 0.017 0.014 1.182 0.700 1.213 0.403 40.3 0.509
Papilio ni eus 0.016 0.012 1.310 0.641 1.299 0.430 43.0 0.023
G aphium philonoe 0.016 0.015 1.090 0.606 0.993 0.457 45.7 0.901
G aphium po haon 0.016 0.013 1.202 0.553 1.087 0.483 48.3 0.660
Danaus ch ysippus 0.015 0.014 1.117 0.276 0.991 0.509 50.9 0.002
Euphaed a neoph on 0.015 0.012 1.261 0.482 1.105 0.534 53.4 0.007
Belenois c eona 0.015 0.015 1.010 0.191 0.974 0.559 55.9 0.001
Papilio cons an inus 0.014 0.015 0.930 0.000 0.836 0.584 58.4 0.001
Ma ia Fungomeli e al.: Bu e ly and ege a ion di e si y in e ac ions in A abuko Sokoke Fo es Kenya316
Species a e age sd a io a a a b Cumsum % o o al dissimila i y P- alue
Nep is sp. 0.014 0.014 1.019 0.884 0.548 0.607 60.7 0.957
Cha axes a anes 0.014 0.012 1.104 0.574 0.871 0.630 63.0 0.027
Nephe onia halassina 0.013 0.012 1.125 0.268 0.872 0.652 65.2 0.345
Ac aea sp. 0.013 0.012 1.058 0.612 0.719 0.674 67.4 0.980
Colo is egina 0.012 0.013 0.980 0.397 0.700 0.694 69.4 0.974
B achys egia s Mixed o es
Phalan a phalan ha 0.026 0.025 1.055 2.143 2.320 0.045 4.5 0.073
Hypolimnas misippus 0.022 0.019 1.189 0.959 1.449 0.082 8.2 0.029
Ca opsilia lo ella 0.022 0.020 1.114 1.234 1.560 0.120 12.0 0.285
Appias epaphia 0.021 0.018 1.195 1.549 2.229 0.155 15.5 0.123
Papilio demodocus 0.019 0.018 1.070 1.106 1.518 0.188 18.8 0.192
G aphium po haon 0.019 0.017 1.105 0.553 1.144 0.220 22.0 0.008
G aphium an heus 0.018 0.017 1.088 1.010 1.087 0.251 25.1 0.285
E onia cleodo a 0.018 0.016 1.147 0.700 1.032 0.281 28.1 0.072
Junonia oenone 0.018 0.015 1.226 0.958 0.934 0.311 31.1 0.795
G aphium philonoe 0.018 0.015 1.175 0.606 1.201 0.341 34.1 0.433
Colo is euippe 0.017 0.014 1.208 0.784 1.142 0.370 37.0 0.262
Papilio ni eus 0.017 0.015 1.144 0.641 1.185 0.399 39.9 0.002
Nep is sp. 0.016 0.015 1.060 0.884 0.926 0.427 42.7 0.347
Nephe onia halassina 0.016 0.015 1.060 0.268 0.940 0.454 45.4 0.001
Ac aea sp. 0.016 0.015 1.075 0.612 0.999 0.481 48.1 0.033
Coeliades o es an 0.016 0.016 1.019 0.628 0.921 0.508 50.8 0.999
Eu ema sp. 0.016 0.020 0.797 0.639 0.669 0.535 53.5 0.332
Belenois hysa 0.015 0.013 1.120 0.265 0.922 0.560 56.0 0.001
Papilio cons an inus 0.015 0.015 1.008 0.000 0.843 0.585 58.5 0.001
Euphaed a neoph on 0.013 0.014 0.990 0.482 0.710 0.608 60.8 0.403
Cha axes a anes 0.013 0.012 1.078 0.574 0.823 0.630 63.0 0.092
Junonia na alica 0.012 0.013 0.953 0.845 1.201 0.651 65.1 0.157
Danaus ch ysippus 0.012 0.013 0.896 0.276 0.668 0.671 67.1 0.424
Colo is auxo 0.012 0.013 0.874 0.427 0.547 0.691 69.1 0.831
Cynome a s Fo es edge
Coeliades o es an 0.030 0.016 1.934 0.668 2.529 0.051 5.1 0.001
Phalan a phalan ha 0.021 0.018 1.162 1.564 2.479 0.087 8.7 0.771
Papilio ni eus 0.020 0.009 2.146 0.000 1.299 0.121 12.1 0.001
Junonia oenone 0.020 0.015 1.317 0.841 1.743 0.154 15.4 0.116
Ca opsilia lo ella 0.020 0.018 1.067 1.646 1.415 0.187 18.7 0.849
Papilio demodocus 0.018 0.014 1.349 0.977 1.910 0.218 21.8 0.422
G aphium philonoe 0.018 0.015 1.197 1.190 0.993 0.247 24.7 0.499
Hypolimnas misippus 0.017 0.014 1.228 1.151 1.738 0.275 27.5 0.972
Colo is ione 0.017 0.014 1.153 0.445 1.185 0.303 30.3 0.004
Appias epaphia 0.017 0.015 1.129 1.828 2.052 0.331 33.1 0.902
Colo is egina 0.016 0.014 1.161 1.122 0.700 0.359 35.9 0.153
Eu ema sp. 0.016 0.015 1.091 0.117 1.176 0.386 38.6 0.209
Colo is euippe 0.016 0.014 1.151 0.569 1.144 0.413 41.3 0.663
G aphium an heus 0.016 0.014 1.126 1.018 1.316 0.440 44.0 0.932
E onia cleodo a 0.015 0.013 1.160 0.785 1.213 0.465 46.5 0.968
Danaus ch ysippus 0.015 0.013 1.134 0.163 0.991 0.490 49.0 0.004
Nep is sp. 0.015 0.014 1.035 1.001 0.548 0.514 51.4 0.870
G aphium po haon 0.014 0.013 1.151 0.665 1.087 0.538 53.8 0.931
Euphaed a neoph on 0.014 0.011 1.309 0.379 1.105 0.562 56.2 0.097
Belenois c eona 0.014 0.014 0.994 0.000 0.974 0.586 58.6 0.001
Cha axes a anes 0.014 0.012 1.129 0.000 0.871 0.609 60.9 0.016
Papilio cons an inus 0.013 0.014 0.979 0.000 0.836 0.632 63.2 0.003
Ac aea sp. 0.013 0.012 1.102 0.729 0.719 0.654 65.4 0.982
Hypolimnas decep o 0.013 0.012 1.014 0.741 0.503 0.675 67.5 0.114
Nephe onia halassina 0.012 0.011 1.141 0.465 0.872 0.695 69.5 0.777
Con ibu ions o En omology 75 (2) 2025, 299–318 317
Species a e age sd a io a a a b Cumsum % o o al dissimila i y P- alue
Cynome a s Mixed o es
Phalan a phalan ha 0.026 0.019 1.330 1.564 2.320 0.045 4.5 0.112
Ca opsilia lo ella 0.021 0.018 1.170 1.646 1.560 0.080 8.0 0.604
Papilio ni eus 0.020 0.012 1.649 0.000 1.185 0.115 11.5 0.001
Hypolimnas misippus 0.020 0.017 1.159 1.151 1.449 0.149 14.9 0.425
Colo is egina 0.019 0.017 1.073 1.122 0.429 0.181 18.1 0.003
Appias epaphia 0.018 0.014 1.259 1.828 2.229 0.212 21.2 0.687
G aphium po haon 0.018 0.016 1.154 0.665 1.144 0.243 24.3 0.048
G aphium philonoe 0.017 0.015 1.162 1.190 1.201 0.273 27.3 0.636
Papilio demodocus 0.017 0.014 1.224 0.977 1.518 0.302 30.2 0.832
E onia cleodo a 0.017 0.014 1.169 0.785 1.032 0.330 33.0 0.417
Nep is sp. 0.017 0.015 1.124 1.001 0.926 0.359 35.9 0.267
Junonia oenone 0.017 0.014 1.182 0.841 0.934 0.387 38.7 0.990
Colo is euippe 0.017 0.013 1.263 0.569 1.142 0.416 41.6 0.492
G aphium an heus 0.017 0.015 1.090 1.018 1.087 0.444 44.4 0.857
Ac aea sp 0.015 0.014 1.101 0.729 0.999 0.470 47.0 0.214
Coeliades o es an 0.015 0.014 1.062 0.668 0.921 0.496 49.6 1.000
Nephe onia halassina 0.015 0.014 1.080 0.465 0.940 0.521 52.1 0.023
Belenois hysa 0.014 0.012 1.153 0.444 0.922 0.545 54.5 0.002
Papilio cons an inus 0.014 0.014 1.025 0.000 0.843 0.569 56.9 0.001
Hypolimnas decep o 0.014 0.014 1.018 0.741 0.513 0.593 59.3 0.009
Cha axes a anes 0.014 0.011 1.282 0.000 0.823 0.616 61.6 0.026
Colo is auxo 0.013 0.012 1.092 0.739 0.547 0.639 63.9 0.202
Bicyclussa i za 0.013 0.013 0.951 0.616 0.447 0.661 66.1 0.015
Cha axes candiope 0.012 0.012 1.008 0.705 0.179 0.682 68.2 0.001
Euphaed a neoph on 0.012 0.012 0.997 0.379 0.710 0.703 70.3 0.912
Fo es edge s Mixed o es
Coeliades o es an 0.025 0.015 1.612 2.529 0.921 0.047 4.7 0.003
Junonia oenone 0.018 0.014 1.352 1.743 0.934 0.082 8.2 0.648
Phalan a phalan ha 0.017 0.019 0.901 2.479 2.320 0.114 11.4 0.995
Ca opsilia lo ella 0.017 0.015 1.070 1.415 1.560 0.145 14.5 0.998
Eu ema sp. 0.015 0.013 1.116 1.176 0.669 0.174 17.4 0.471
Colo is ione 0.015 0.014 1.084 1.185 0.662 0.201 20.1 0.189
G aphium philonoe 0.015 0.012 1.231 0.993 1.201 0.229 22.9 0.995
Colo is euippe 0.014 0.011 1.250 1.144 1.142 0.256 25.6 0.994
E onia cleodo a 0.014 0.012 1.185 1.213 1.032 0.283 28.3 0.995
Hypolimnas misippus 0.014 0.013 1.091 1.738 1.449 0.310 31.0 1.000
G aphium an heus 0.014 0.013 1.103 1.316 1.087 0.336 33.6 0.999
G aphium po haon 0.013 0.012 1.115 1.087 1.144 0.361 36.1 0.999
Papilio demodocus 0.013 0.013 1.005 1.910 1.518 0.387 38.7 0.997
Danaus ch ysippus 0.013 0.012 1.097 0.991 0.668 0.411 41.1 0.143
Appias epaphia 0.013 0.011 1.117 2.052 2.229 0.434 43.4 1.000
Ac aea sp. 0.013 0.011 1.104 0.719 0.999 0.458 45.8 0.991
Belenois c eona 0.012 0.013 0.995 0.974 0.175 0.482 48.2 0.001
Euphaed a neoph on 0.012 0.010 1.154 1.105 0.710 0.504 50.4 0.948
Papilio cons an inus 0.012 0.011 1.054 0.836 0.843 0.526 52.6 0.087
Nephe onia halassina 0.012 0.011 1.046 0.872 0.940 0.548 54.8 0.919
Nep is sp. 0.011 0.011 1.056 0.548 0.926 0.570 57.0 1.000
Belenois hysa 0.011 0.010 1.109 0.329 0.922 0.592 59.2 0.520
G aphium colonna 0.011 0.011 0.993 0.771 0.577 0.612 61.2 0.581
Cha axes a anes 0.011 0.009 1.157 0.871 0.823 0.633 63.3 0.960
Colo is egina 0.011 0.011 0.964 0.700 0.429 0.653 65.3 0.998
Junonia na alica 0.010 0.011 0.912 0.992 1.201 0.671 67.1 0.929
Melani is leda 0.010 0.009 1.035 0.648 0.574 0.689 68.9 0.174
Hypolimnas decep o 0.010 0.011 0.859 0.503 0.513 0.707 70.7 0.882
Ma ia Fungomeli e al.: Bu e ly and ege a ion di e si y in e ac ions in A abuko Sokoke Fo es Kenya318
Supplemen a y ma e ial 1
Suppl. igu es S1–S3
Au ho s: Ma ia Fungomeli, Ma in Wieme s, Lucia
Calde ini, Alessand o Chia ucci
Da a ype: pd
Explana ion no e: igu e S1. F equency anking o bu -
e ly species abundance wi hin he o es ypes o
B achys egia, Cynome a, Mixed o es and Fo es
edge in A abuko Sokoke Fo es , Kenya. igu e S2.
Abundance p opo ions o bu e ly species wi h mono-
phagous, oligophagous and polyphagous la al eed-
ing habi s ac oss he ou o es ypes o B achys egia,
Cynome a, Mixed o es and Fo es edge in A abuko
Sokoke Fo es , Kenya. igu e S3. Bu e lies in A abu-
ko Sokoke Fo es , in he mixed o es ege a ion ype,
eeding om elephan dung du ing he ield sampling
in he d y season. Pho o c edi s: Ma ia Fungomeli.
Copy igh no ice: This da ase is made a ailable unde
he Open Da abase License (h p://openda acommons.
o g/licenses/odbl/1.0). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow us-
e s o eely sha e, modi y, and use his Da ase while
main aining his same eedom o o he s, p o ided
ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/con ib.en omol.75.e155016.
suppl1
