73
Changes in species and phylogene ic di e si y in opical
seasonal ain o es s on s eep ka s hillslopes in sou hwes e n
China: implica ions o conse a ion
Qingling Pang1* , Cong Hu1* , Chaohao Xu1, Chao ang Zhong1, Zhonghua Zhang1,2,3 , Gang Hu1,2,3
1 Key Labo a o y o En i onmen Change and Resou ces Use in Beibu Gul , Minis y o Educa ion, Nanning No mal Uni e si y, Nanning 530100, China
2 Guangxi Key Labo a o y o Ea h Su ace P ocesses and In elligen Simula ion, Nanning No mal Uni e si y, Nanning 530100, China
3 Key Labo a o y o Wildli e E olu ion and Conse a ion in Moun ain Ecosys em o Guangxi, Nanning No mal Uni e si y, Nanning 530100, China
Co esponding au ho s: Zhonghua Zhang ([email p o ec ed]); Gang Hu ([email p o ec ed])
Copy igh : © Qingling Pang e al.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
T opical ka s seasonal ain o es s (TKSR) in sou hwes e n China ha e high le els o
biodi e si y and a signi ican numbe o endemic species. Howe e , unde s anding he
dis ibu ion o plan di e si y and he mechanisms d i ing communi y assembly in hese
di e se and he e ogeneous ka s o es s emains limi ed. This s udy examined he spe-
cies and phylogene ic di e si y o he he b, sh ub, and ee laye s in he TKSR along
s eep hillslopes wi hin ka s peak-clus e dep essions in sou hwes e n China. Species
ichness in he he b, sh ub, and ee laye s showed an in e ed U-shape pa e n, wi h
ichness lowe in he middle and highe on he sides o he hillslope. The uppe slope
had a highe le el o species ichness han he dep essions and he lowe and middle
slopes. The phylogene ic s uc u es o he he b and ee laye s ended o be clus e ed,
whe eas he sh ub laye exhibi ed a di e gen phylogene ic s uc u e. These indings
indica e ha communi y assembly in he TKSR is in luenced by habi a il e ing and
compe i i e exclusion. Soil o al phospho us (STP) and soil a ailable phospho us (SAP)
we e iden i ied as signi ican ac o s a ec ing species di e si y ac oss he h ee laye s,
whe eas he ock ou c op a e was a signi ican ac o a ec ing he phylogene ic s uc-
u e. In he sh ub and ee laye s, STP and SAP we e impo an de e minan s o phylo-
gene ic di e si y. These esul s highligh he impac o en i onmen al he e ogenei y on
species and phylogene ic di e si y in he TKSR. Fu he mo e, dep essions and uppe
slopes wi h high species and phylogene ic di e si y we e iden i ied as p io i y a eas o
conse a ion in he ka s peak-clus e dep essions o sou hwes e n China.
Key wo ds: Communi y assembly, ka s o es , phylogene ic s uc u e, plan di e si y,
slope posi ion
In oduc ion
Changes in plan di e si y along en i onmen al g adien s a e impo an in he ield
o ege a ion ecology (Liu 2017). Slope, as a opog aphical ac o , plays a c ucial
ole in con olling he spa ial dis ibu ion o wa e , ligh , hea , and soil nu ien s, and
in luences he abundance, dis ibu ion, and di e si y o ege a ion in moun ainous
Academic edi o : Ilona Blino a
Recei ed:
19 June 2025
Accep ed:
12 Oc obe 2025
Published:
4 No embe 2025
ZooBank: h ps://zoobank.o g/
FDBA1FC3-45CC-4410-8568-
B24703AF42A6
Ci a ion: Pang Q, Hu C, Xu C, Zhong C,
Zhang Z, Hu G (2025) Changes in species
and phylogene ic di e si y in opical
seasonal ain o es s on s eep ka s hillslopes
in sou hwes e n China: implica ions
o conse a ion. Na u e Conse a ion
60: 73–97. h ps://doi.o g/10.3897/
na u econse a ion.60.162595
Na u e Conse a ion 60: 73–97 (2025)
DOI: 10.3897/na u econse a ion.60.162595
* These au ho s con ibu ed equally o his wo k.
74
Na u e Conse a ion 60: 73–97 (2025), DOI: 10.3897/na u econse a ion.60.162595
Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
and hilly a eas (Zeng e al. 2014). Plan di e si y pa e ns along slopes o en ex-
hibi complex and non-mono onic changes ha a e closely ela ed o opog aphy,
wa e , and hea condi ions, as well as a ious li e o ms, including ees, sh ubs,
and he bs (Deák e al. 2021; Sha ma and Kala 2022). Many s udies ha e in es i-
ga ed he impac o slope on ege a ion in a ious ecosys em ypes (Zeng e al.
2014; Be y e al. 2016; Zhang e al. 2020; Dong e al. 2022; Liang e al. 2024). Un-
de s anding plan di e si y pa e ns along slopes and he unde lying mechanisms
is c ucial o elucida ing he ela ionship be ween ege a ion and he en i onmen .
Species di e si y is a cen al aspec o biodi e si y and ecology esea ch,
e lec ing communi y s uc u e, de elopmen al s ages, s abili y, and habi a di -
e ences, and can also e eal he o ganiza ional le els o plan communi ies (Li
e al. 2019). Unde s anding he in luence o en i onmen al ac o s on species
di e si y is an impo an ecological ques ion (Weigel e al. 2019). Plan commu-
ni ies a e ypically composed o species wi h a ious li e his o ies ha occupy
dis inc ecological niches (Ne esheim e al. 2018). As he communi y en i-
onmen changes, he niches occupied by hese species can change, leading
o di e en pa e ns o he e ogenei y in he communi y (Baga ia e al. 2019).
Consequen ly, di e en communi ies o en exhibi di e ences in species com-
posi ion, s uc u e, and unc ional ai s; elucida ing hese di e ences is c ucial
o unde s anding how species di e si y a ies ac oss di e en en i onmen s
(Geng e al. 2022). Phylogene ic di e si y can be used o assess he uniqueness
o biological axa in speci ic spa io empo al con ex s based on he e olu iona y
his o y o he species. I is also indica i e o biodi e si y conse a ion because
i can be used o iden i y phylogene ic ela ionships and e olu iona y in o ma-
ion among di e en species (Chen e al. 2021). Phylogene ic s uc u es com-
plemen phylogene ic di e si y and can be used o in e he ecological p ocess-
es a ec ing communi y assembly. When in eg a ed, he composi ion, o igin,
and e olu iona y p ocesses o species can be explo ed om bo h e olu iona y
and ecological pe spec i es, he eby p omo ing he a ional conse a ion o
plan di e si y (Ci and Li 2017). Phylogene ic di e si y in e p e s a species’ e o-
lu iona y his o y om a phylogene ic ela ionship pe spec i e. When combined
wi h species di e si y, i g ea ly enhances ou unde s anding o popula ion dy-
namics and ecological niche di e en ia ion (Zeng e al. 2014), p o iding a new
pe spec i e o biodi e si y conse a ion. Al hough p e ious s udies ha e o-
cused on species pa e ns and phylogene ic di e si y in di e en o es ypes
wo ldwide (Mye s e al. 2013; Qian e al. 2014; Yang e al. 2014; Guo e al. 2018;
Gas aue e al. 2020), unde s anding o ege a ion dis ibu ion cha ac e is ics
in special land o ms, such as ka s landscapes, emains limi ed.
Ka s landscapes a e o med by he e ec s o ain all and g oundwa e on
ca bona e bed ock (De Waele 2017). These landscapes a e widesp ead, occu -
ing on app oxima ely 15%–20% o he Ea h’s ice- ee land su ace, wi h he
la ges con inuous a ea spanning oughly 0.51 million km2 in sou hwes e n Chi-
na (Jiang e al. 2014). Owing o hei high sensi i i y and ulne abili y, he con-
se a ion o biodi e si y in ka s a eas is o g ea impo ance (Sun e al. 2020).
Ka s a eas in sou hwes e n China a e ecognized as one o he 25 global bio-
di e si y ho spo s because o hei unique biomes. The dis inc i e land o ms
hey c ea e consis o peak-clus e dep essions, cha ac e ized as closed and
la dep essions su ounded by a se ies o hills. Ka s peak-clus e dep essions
con ibu e o he complexi y and di e si y o land o ms in ka s a eas (Yang e
75
Na u e Conse a ion 60: 73–97 (2025), DOI: 10.3897/na u econse a ion.60.162595
Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
al. 2014) and display apidly changing opog aphical cha ac e is ics, ansi ion-
ing om cli s o dep essions o e sho ho izon al dis ances. In addi ion, he
po en ial ene gy dec eases d ama ically om he op slope o he dep ession
a ea, esul ing in a signi ican po en ial g adien . These peak-clus e dep es-
sions p o ide he opog aphic and hyd ological condi ions necessa y o he de-
elopmen o a ca ena y pa e n o soil and ege a ion (Meng e al. 2022). Ka s
peak-clus e dep essions in sou hwes e n China a e gene ally a low ele a ions
(be ween 200 and 500 m) bu o en exhibi s eep slopes (g ea e han 30°). In
hese a eas, ele a ion is no conside ed he p ima y ac o in luencing species
di e si y in communi ies, especially in low- and small-scale egions (Zeng e
al. 2014). Slope posi ion has been ecognized as one o he majo opog aphic
ac o s ha de e mine mic oclima es and ecological niches o species in ka s
moun ain egions, as i in luences a ious aspec s such as sunligh , soil dep h,
soil mois u e, and nu ien a ailabili y (Gong e al. 2008; Wein aub e al. 2015).
Moun ainous and hilly a eas encompassing s eep en i onmen al g adien s
in small geog aphic a eas a e ou s anding na u al labo a o ies o biodi e si y
s udies because nume ous en i onmen al ac o s, such as empe a u e (Wu e
al. 2013), p oduc i i y (Ramí ez-Bau is a and Williams 2019), and an h opogen-
ic dis u bance (San illán e al. 2020), can co ela e wi h ele a ion and conse-
quen ly exe e ec s on biological communi ies. Along slope posi ion g adien s,
ka s peak-clus e dep essions exhibi highly he e ogeneous mic ohabi a s due
o a ia ions in mul iple en i onmen al ac o s, esul ing in signi ican di e enc-
es in biodi e si y. Fo example, Peng e al. (2019) epo ed ha slope was he
majo d i e o he spa ial dis ibu ion o soil mic obial ichness and di e si y
in a ka s o es due o i s e ec s on plan cha ac e is ics (i.e., ee Shannon
di e si y and ee densi y) and soil p ope ies (i.e., soil pH and a ailable phos-
pho us). Simila ly, Sun e al. (2020) e ealed spa ial incong uence among he
axonomic, unc ional, and phylogene ic di e si y o small mammals along ele-
a ion g adien s in ka s moun ains. O he s udies ha e ocused on a ia ions
in soil physicochemical p ope ies and plan –li e –soil ecological s oichiome-
y along slope posi ion g adien s in ka s peak-clus e dep essions (Zeng e al.
2015; Chen e al. 2019; Hu e al. 2020). Howe e , i emains unclea how plan
species and phylogene ic di e si y change along slope posi ion g adien s in
ka s peak-clus e dep essions and he d i ing ac o s behind hese changes.
The Nonggang Na ional Na u e Rese e in sou hwes e n China is a ep esen-
a i e o ka s peak-clus e dep ession land o ms wo ldwide. The opical ka s
seasonal ain o es s (TKSR) in his ese e a e cha ac e ized by complex s uc-
u es, ich species composi ion, and p ominen endemic elemen s. This a ea is
also one o he 14 key a eas o e es ial biodi e si y o in e na ional signi icance
in China (Huang e al. 2013). Thus, he TKSR is i eplaceable and signi ican o
he es o a ion and p o ec ion o ka s o es ecosys ems (Zeng e al. 2023). Re-
cen ecological s udies on he TKSR ha e ocused on species composi ion, spa ial
dis ibu ion, and di e si y (Ma e al. 2023). S udies ha e ound ha he di e si y
o woody plan species in he TKSR is unimodal, bu he e is also a bimodal pa -
e n ha inc eases wi h ele a ion, and he di e si y pa e n is s ongly in luenced
by opog aphic he e ogenei y (Huang e al. 2016). Fu he mo e, Guo e al. (2017,
2018) in es iga ed he e ec s o de e minis ic and s ochas ic p ocesses on com-
muni y assembly in he TKSR a a 15-ha plo scale and ound ha bo h p ocesses
play impo an oles. Al hough i is ecognized ha opog aphy in luences he spa-
76
Na u e Conse a ion 60: 73–97 (2025), DOI: 10.3897/na u econse a ion.60.162595
Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
ial dis ibu ion o ees in he TKSR, he dominan bio ic and abio ic ac o s ha a -
ec he dis ibu ion pa e ns o species and phylogene ic di e si y emain unclea .
In his s udy, plo su eys and en i onmen al ac o measu emen s we e
conduc ed in he TKSR o he Nonggang Na ional Na u e Rese e wi h a ocus
on di e en slopes (dep ession, lowe slope, middle slope, and uppe slope).
G adien changes in communi y species and phylogene ic di e si y ac oss
h ee o es s a a (he b, sh ub, and ee laye s) and he ac o s in luencing
hese changes we e examined. The objec i e o his s udy was o e eal (1) he
spa ial pa e ns o species di e si y, phylogene ic di e si y, and s uc u e wi hin
slope g adien s in he he b, sh ub, and ee laye s and (2) he p ima y en i on-
men al ac o s in luencing hese spa ial pa e ns. The esul s p o ide a scien-
i ic basis o imp o ing unde s anding o o es communi y s uc u e, ecosys-
em unc ions, and ecological conse a ion in ka s peak-clus e dep essions.
Ma e ials and me hods
S udy a ea
The Nonggang Na ional Na u e Rese e, loca ed in he sou he n Guangxi Zhuang
Au onomous Region, sou hwes e n China (22°13'56"–22°39'09"N, 106°42'28"–
107°04'54"E; Fig. 1), co e s an a ea o 10,080 ha. The egion has a opical mon-
soon clima e, wi h an a e age annual empe a u e o app oxima ely 22 °C. The
empe a u e anges om abo e 13 °C in Janua y, he coldes mon h, o abo e
28 °C in July, he ho es mon h. The annual accumula ed empe a u e anges om
7,400 o 7,800 °C. Annual a e age ain all anges om 1,200 o 1,500 mm and is
concen a ed be ween May and Sep embe . The bed ock o he a ea is p ima ily
Figu e 1. Geog aphical loca ion o he s udy a ea, wi h pho og aphs showing opical ka s seasonal ain o es s along
s eep ka s hillslopes.
77
Na u e Conse a ion 60: 73–97 (2025), DOI: 10.3897/na u econse a ion.60.162595
Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
composed o limes one, and he main soil ypes a e p imi i e calca eous, b own
calca eous, and black calca eous. The opog aphy o he egion is cha ac e ized
by ypical ka s peak-clus e dep ession land o ms, wi h dep essions o med by
clus e ed peaks sha ing a common base and a unnel-shaped landscape. Ele a-
ion anges om 150 o 600 m. Unique land o ms, such as peak clus e s, o es s,
dep essions, and unnels, esul in signi ican a ia ions in ligh a ailabili y, soil
hickness, and mois u e. O he mic o-land o m ea u es o med by nume ous ock
ou c ops—such as s one gullies, s one su aces, and dep essions—con ibu e o
he he e ogenei y o small-scale habi a s. The ese e con ains 1,752 eco ded
species o ascula plan s belonging o 810 gene a and 184 amilies (Huang e al.
2013). Among hese, 33 species a e a e and endange ed in China, and 278 species
a e endemic o ka s habi a s (Wang e al. 2014). The dominan ee species in he
TKSR include Excen odend on onkinense (A. Che .) H.T. Chang & R.H. Miao, Ca-
mellia pe elo ii (Me .) Sealy, Cephalomappa sinensis (Chun & F.C. How) Kos e m.,
Ga cinia paucine is Chun ex F.C. How, and Deu zian hus onkinensis Gagnep.
Da a collec ion
Du ing he peak g owing season om July o Sep embe 2020, o es plo s we e
es ablished a di e en slope posi ions based on ele a ion a ia ions in he ka s
peak-clus e dep essions in he s udy a ea (Fig. 1). The e we e 10 plo s, each
wi h an a ea o 20 × 20 m, a ou slope posi ions—dep ession (DE), lowe slope
(LS), middle slope (MS), and uppe slope (US)—wi h a spacing o mo e han 30 m
be ween plo s. A o al o 40 plo s we e es ablished along s eep ka s hillslopes
in he s udy a ea. T ee-laye in es iga ions included species names, diame e s
a b eas heigh (DBH), and heigh s. Wi hin each plo , wo 5 × 5 m subplo s we e
andomly es ablished o in es iga e he sh ub laye , including species names, a -
e age heigh s, co e ages, and numbe s. Addi ionally, ou 1 × 1 m subplo s we e
andomly es ablished o in es iga e he he b laye , including species names,
numbe s, heigh s, and co e ages. All ascula plan s in each plo and subplo
we e in en o ied, wi h species iden i ica ions e i ied by wo plan axonomy ex-
pe s. Fu he mo e, la i ude and longi ude coo dina es, ele a ion, aspec , slope
posi ion, and ock ou c op a e we e eco ded. Th ee expe ienced obse e s i-
sually es ima ed he slope posi ion and ock ou c op a e, and a handheld GPS
(Ga min GPSMAP 60CSx) was used o all o he measu emen s. Slope posi ions
we e ep esen ed by ca ego ical a iables 1, 2, 3, and 4, co esponding o dep es-
sion, lowe slope, middle slope, and uppe slope, espec i ely (Qiu and Zhang
2000). The slope aspec was con e ed om a ange o 0–360° azimu h o a al-
ue anging om 0 o 1 using a p e iously documen ed o mula (Yu e al. 2013).
Wi hin each plo , soil samples we e collec ed a dep hs o 0–20 cm using he
i e-poin sampling me hod (Lu e al. 2023). The collec ed soil samples we e
mixed and e u ned o he labo a o y. A e ai -d ying and sie ing, he soil samples
we e subjec ed o chemical analyses o de e mine a ious soil pa ame e s. The
measu ed indica o s included soil pH, exchangeable calcium (ECa), exchange-
able magnesium (EMg), a ailable phospho us (AP), a ailable po assium (AK),
ammonium ni ogen (NH4
+-N), ni a e ni ogen (NO3
--N), o ganic ma e (SOM),
o al ni ogen (TN), o al phospho us (TP), and o al po assium (TK). Soil pH was
de e mined in a suspension wi h a 1:2.5 soil- o-wa e a io. ECa and EMg we e
measu ed using a omic abso p ion spec opho ome y (AA-7000, Shimadzu,
78
Na u e Conse a ion 60: 73–97 (2025), DOI: 10.3897/na u econse a ion.60.162595
Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
Kyo o, Japan). NH4
+-N was quan i ied using he indophenol blue colo ime ic
me hod, and NO3
--N was measu ed using he cadmium educ ion me hod ol-
lowed by a sul anilamide–NAD eac ion. SOM con en was de e mined using he
adi ional po assium dich oma e oxida ion me hod. TN was measu ed using he
semi-mic o Kjeldahl me hod. AP and TP le els we e de e mined using he molyb-
denum–an imony colo ime ic me hod. AK and TK le els we e measu ed using
he sodium hyd oxide usion lame pho ome y me hod. The su ace soil wa e
con en (SWC) o each plo was measu ed using a po able h ee-pa ame e soil
es e (WET-2, Del a-T De ices L d., Uni ed Kingdom) ollowing he i e-poin
sampling me hod (Lu e al. 2023). These me hods o measu ing soil p ope ies
we e desc ibed in de ail by Lu (1999) and Bao (2000). En i onmen al a iables
(mean ± SD) in ou slope posi ions a e p esen ed in Suppl. ma e ial 1: able S1.
Da a analysis
Species di e si y
The plan species names, along wi h hei amily and genus in o ma ion wi hin
he s udy plo s, we e e i ied acco ding o he APG III sys em using he plan -
lis package (Zhang 2018). Species ichness (S), Simpson index (D), Shannon–
Wiene index (H), and Pielou index (J) we e calcula ed o assess species di-
e si y in he plan communi ies (MacDonald e al. 2017). These indices we e
compu ed using he egan package in R e sion 4.2.2 (Oksanen e al. 2019).
The o mulae used o he calcula ions a e as ollows:
(1)
(2)
J = H / lnS (3)
whe e S ep esen s he species ichness wi hin he communi y plo and Pi ep-
esen s he impo ance alue o species i in he communi y.
Phylogene ic s uc u e and di e si y
A phylogene ic ee o ascula plan s was cons uc ed using he V.PhyloMake
package (Jin and Qian 2019), inco po a ing e olu iona y b anch leng hs based
on he “GBOTB.ex ended. e” amewo k ollowing he me hodological ap-
p oach o scena io S3. This scena io employs a e age dis ances o cons ain
phylogene ic ip posi ions, he eby educing bias in oduced by poly omies. The
“GBOTB.ex ended. e” ee in eg a es he e olu iona y b anches o seed plan s
om GBOTB (Smi h and B own 2018) and e ns om he upda ed, e ised, and
expanded phylogene ic sys em o Zanne e al. (2014).
The Fai h Di e si y Index (PD) was used o measu e phylogene ic di e si y (Fai h
1992). The ne ela edness index (NRI) and nea es axon index (NTI) we e used
o de e mine he phylogene ic s uc u e. NRI and NTI ep esen he s anda dized
alues o he mean pai wise dis ance (MPD) and mean nea es axon dis ance
79
Na u e Conse a ion 60: 73–97 (2025), DOI: 10.3897/na u econse a ion.60.162595
Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
(MNTD), espec i ely. The NRI p ima ily desc ibes he o e all phylogene ic ela-
ionships among species in a communi y, whe eas he NTI ocuses on phyloge-
ne ic ela ionships a he e minal b anches o he phylogene ic ee. Compa ed
wi h NRI, NTI places g ea e emphasis on desc ibing he phylogene ic dis ances
be ween neighbo ing species (Webb e al. 2002). MPD and MNTD we e calcula -
ed using he comdis and comdis n unc ions in he pican e package (Kembel e
al. 2008). The ac ual alues o each communi y we e compa ed using an uncon-
s ained null model (Kembel and Hubbell 2006), in which he e minal b anches
o he phylogene ic ee we e andomly pe mu ed 999 imes o all species in he
communi y. The di e ences we e hen calcula ed, and he NRI and NTI alues o
each communi y we e ob ained (Webb e al. 2002; Swenson e al. 2007).
PD = ∑Lb (4)
(5)
(6)
whe e Lb ep esen s he b anch leng h connec ing he species on he ee. MP-
Dobse ed and MNTDobse ed a e he obse ed alues o he a e age pai wise
phylogene ic dis ance and mean nea es axon dis ance, espec i ely, o each
plo . MPD andomized and MNTD andomized a e he a e age pai wise phylo-
gene ic dis ance and mean nea es axon dis ance, espec i ely, ob ained by
andomly gene a ing null models o he communi y. sdMPD andomized and
sdMNTD andomized a e he s anda d de ia ions o he a e age pai wise phylo-
gene ic dis ance and mean nea es axon dis ance ob ained om he andomly
gene a ed null models o he communi y, espec i ely. I he NRI (o NTI) > 0, he
phylogene ic ela ionships among indi iduals in he communi y a e close han
hose expec ed om a andomly gene a ed null model, sugges ing phylogene ic
clus e ing. Con e sely, i he NRI (o NTI) < 0, i sugges s phylogene ic o e dis-
pe sion, indica ing ha he phylogene ic ela ionships among indi iduals in he
communi y a e mo e di e gen han expec ed om he null model (Webb e al.
2002). I he NRI (o NTI) < −1.96, i indica es signi ican phylogene ic e enness
compa ed wi h he null model, whe eas an NRI (o NTI) > 1.96 indica es signi i-
can phylogene ic clus e ing compa ed wi h he null model (Vamosi e al. 2009).
S a is ical analysis
Assuming ha he da a we e no mally dis ibu ed and sa is ied he assump ion
o homogenei y o a iances, one-way ANOVA was used o compa e di e enc-
es in en i onmen al a iables among he di e en slope posi ions. Fo mul iple
compa isons o plan di e si y be ween slope posi ions, he leas signi ican
di e ence (LSD) me hod was applied i a iances we e equal; howe e , i a i-
ances we e no equal, he Games–Howell es was used. Pea son co ela ion
analysis was conduc ed o examine ela ionships be ween species, phylogene -
ic di e si y, and en i onmen al a iables. Fac o s wi h > 0.6 we e emo ed. The
emaining ac o s we e hen included in mul iple s epwise eg ession analyses,
along wi h di e si y indices and phylogene ic s uc u es, o iden i y he main
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Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
ac o s in luencing species and phylogene ic di e si y. The s epwise eg ession
p ocedu e was based on he Akaike In o ma ion C i e ion (AIC), in which a i-
ables we e added o emo ed by selec ing he model wi h he minimum AIC
alue. Residual analysis con i med ha he model sa is ied he assump ions
o no mali y, linea i y, and homoscedas ici y, he eby alida ing he use o a e-
g ession model based on he Gaussian dis ibu ion. The ac o s e ained in he
eg ession equa ions we e quan i ied using he dacca.hp package (Lai e al.
2022) in R 4.2.2 o de e mine hei explana o y powe o he di e si y indices
h ough hie a chical pa i ioning analysis. One-way ANOVA and Pea son co -
ela ion analysis we e pe o med using SPSS 26.0 (SPSS Inc., Chicago, IL, USA).
Resul s
Species and phylogene ic di e si y a di e en slopes
Species di e si y in he he b, sh ub, and ee laye s ini ially dec eased and hen in-
c eased wi h slope posi ion (Fig. 2). In he he b laye , species ichness in he mid-
dle slope was signi ican ly lowe han ha in he lowe and uppe slopes (p < 0.05),
and he Pielou index showed an inc easing end, wi h alues o bo h he middle
and uppe slopes signi ican ly highe han hose in he lowe slopes (p < 0.05).
The e we e no signi ican di e ences in species di e si y in he sh ub laye among
he di e en slope posi ions. In he ee laye , species ichness ini ially dec eased
and hen inc eased wi h slope posi ion, whe eas he o he indices showed an o e -
all inc easing end, wi h no signi ican di e ences among slope posi ions.
Figu e 2. Changes in species di e si y a di e en slope posi ions. Di e en lowe case le e s indica e a signi ican di -
e ence be ween communi ies a di e en slope posi ions (p < 0.05). DE, dep ession; LS, lowe slope; MS, middle slope;
US, uppe slope.
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Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
Phylogene ic di e si y in he he b, sh ub, and ee laye s ini ially dec eased
and hen inc eased wi h slope posi ion (Fig. 3). Speci ically, phylogene ic di-
e si y in he lowe and uppe slopes was signi ican ly highe han ha in he
middle slope in he he b laye (p < 0.05). In con as , he e we e no signi ican
di e ences in phylogene ic di e si y among slope posi ions in he sh ub and
ee laye s. Ac oss he h ee laye s, phylogene ic di e si y was highes in he
ee laye , ollowed by he he b and sh ub laye s.
The NRI and NTI in he he b, sh ub, and ee laye s showed inconsis en
pa e ns in communi y species phylogene ic ela ionships ac oss he slope
posi ions (Fig. 4). In he he b laye , bo h NRI and NTI indica ed phylogene ic
clus e ing ac oss he ou slope posi ions. In he sh ub laye , bo h NRI and NTI
exhibi ed phylogene ic di e gence on he lowe and uppe slopes, whe eas
he middle and dep ession slopes showed a coexis ence o phylogene ic di-
e gence and clus e ing, wi h signi ican di e ences in NRI among he slope
posi ions (p < 0.05). In he ee laye , bo h NRI and NTI exhibi ed an o e all
pa e n o phylogene ic clus e ing on he lowe and uppe slopes. In con as ,
he dep ession and middle slopes showed a coexis ence o phylogene ic di-
e gence and clus e ing. O e all, he he b and ee laye s showed phylogene ic
clus e ing, whe eas he sh ub laye exhibi ed phylogene ic di e gence.
Species and phylogene ic di e si y in ela ion o en i onmen al a iables
Co ela ion analysis e ealed ha species di e si y in he he b laye was signi i-
can ly and nega i ely co ela ed wi h NO3
--N, SOM, and SWC, and signi ican ly
and posi i ely co ela ed wi h SP, SA, and SD (p < 0.05). The phylogene ic s uc-
u e was signi ican ly and posi i ely co ela ed wi h AK, TP, and BA (p < 0.05).
Phylogene ic di e si y was signi ican ly and nega i ely co ela ed wi h NO3
--N
and SOM con en s (p < 0.05; Fig. 5).
In he sh ub laye , species di e si y was signi ican ly and nega i ely co ela -
ed wi h TP and BA and signi ican ly and posi i ely co ela ed wi h SA and pH
Figu e 3. Phylogene ic di e si y a di e en slope posi ions. Di e en lowe case le e s
indica e a signi ican di e ence be ween communi ies a he a ious slope posi ions
(p < 0.05). DE, dep ession; LS, lowe slope; MS, middle slope; US, uppe slope.
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Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
s uc u al pa e ns wi hin plan communi ies, which a e in luenced by mul iple en-
i onmen al ac o s a di e en le els. Speci ically, phylogene ic agg ega ion in he
he b laye was associa ed wi h inc eases in soil AK, TP, and BA. In he sh ub laye ,
agg ega ion was d i en by ele a ed soil NH4
+-N, NO3
--N, pH, and BA. Con e sely,
he ee laye esponded o inc eases in soil ECa, EMg, TK, TN, TP, SOM, and ROR.
Howe e , an inc ease in SD p omo ed an o e all agg ega ion end in he phylo-
gene ic ela ionships o ee laye species, whe eas SOM had he opposi e e ec ,
inc easing he phylogene ic dis ance be ween adjacen species and a ec ing he
phylogene ic s uc u e o he ee laye . The phylogene ic s uc u e o plan com-
muni ies is closely ela ed o he esponse o plan s o hei en i onmen , which in
u n a ec s he ecological p ocesses ha domina e communi y assembly.
The phylogene ic s uc u es o he he b, sh ub, and ee laye s exhibi ed in-
consis en pa e ns along he slope g adien . The he b laye s ac oss he di -
e en slopes exhibi ed agg ega ed phylogene ic s uc u es. In addi ion o he
in luence o mic ohabi a condi ions, he phylogene ic s uc u e o he he b lay-
e may be ela ed o he li e his o ies o he baceous plan s (Qian e al. 2014).
He baceous plan s ha e sho li e cycles and s ong dispe sal abili ies, can ap-
idly occupy sui able habi a s, exhibi la ge popula ion luc ua ions, and easily
o m agg ega ed phylogene ic s uc u es (Niu e al. 2011). The dispe sal o ee
seeds may also in luence he phylogene ic s uc u e o he he b laye communi-
y. He b laye s include woody plan seedlings, and he limi ed dispe sal o seeds
om pa en ees in he o es leads o a clus e ed pa e n o seedlings, esul -
ing in an agg ega ed phylogene ic s uc u e in he he b laye communi y (Niu e
al. 2011). The phylogene ic s uc u e o he sh ub laye exhibi ed di e gence in
he dep ession and uppe slope and agg ega ion in he middle slope communi-
ies. In con as , he lowe slope communi ies showed inconsis en NRI and NTI
alues o he phylogene ic ela ionships o communi y species, wi h NRI indi-
ca ing di e gence and NTI indica ing agg ega ion. In he ee-laye communi ies,
he NRI and NTI o he dep ession and middle slopes exhibi ed inconsis en phy-
logene ic s uc u es. In he dep ession, NRI showed di e gence, whe eas NTI
showed agg ega ion. Con e sely, he opposi e pa e n was obse ed in he mid-
dle slope communi ies, while he phylogene ic s uc u es o he uppe and lowe
slope communi ies we e agg ega ed. In high-al i ude a eas, due o en i onmen-
al s ess, species wi hin communi ies unde go con e gen e olu ion, esul ing
in a end owa d agg ega ed phylogene ic s uc u es (Kluge and Kessle 2011;
You e al. 2013). Simila ly, esea ch has shown ha he phylogene ic s uc u es
o e e g een b oad-lea ed o es communi ies exhibi agg ega ion in low-al i-
ude a eas and di e gence in high-al i ude a eas (Huang e al. 2010). In his
s udy, he ha sh habi a condi ions on he uppe slope may ha e caused s ong
en i onmen al s ess, p omo ing con e gen e olu ion in plan s and esul ing in
agg ega ed phylogene ic s uc u es in he ee-laye communi ies. Con e sely,
compe i ion o limi ed esou ces among plan s may ha e caused he phylo-
gene ic s uc u e o sh ub laye plan s o di e ge. The inconsis en changes in
communi y NRI and NTI alues a e ela ed o hei di e en emphases, indica -
ing ha dis inc ecological p ocesses in luence he assembly o he b, sh ub,
and ee laye communi ies. The di e en ways in which communi ies espond
o he en i onmen a ec he ela i e impo ance o ecological p ocesses in
communi y assembly. The esul s indica e ha de e minis ic p ocesses (en i-
onmen al il e ing and compe i i e exclusion) ha e signi ican impac s on com-
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Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
muni y assembly in he TKSR, while some communi ies may also be in luenced
by neu al p ocesses (NRI o NTI < −1.96) (Vamosi e al. 2009).
Conse a ion o ka s o es s
Ka s landscapes a e c ucial biodi e si y ese oi s, pa icula ly in Asia, hos ing
a weal h o endemic species (Clemen s e al. 2006). The s eep ka s egions o
sou hwes e n China a e ecognized as biodi e si y ho spo s, no able o hei
ema kable plan species ichness and signi ican conse a ion alue (Guo e
al. 2018; Geekiyanage e al. 2019). P o ec ing biodi e si y and ehabili a ing
deg aded ege a ion in hese a eas emain pi o al ecological p io i ies o he
Chinese go e nmen and esea che s (Chen e al. 2024). Conse a ion s a -
egies should inco po a e phylogene ic di e si y in o assessmen amewo ks,
complemen ing adi ional species di e si y me ics o e lec e olu iona y ela-
ionships (Tie je e al. 2023). Consequen ly, a eas wi h high and unique phyloge-
ne ic di e si y should be p io i ized o conse a ion e o s (Zhang e al. 2024).
Ou s udy demons a es ha dep essions and uppe slopes exhibi g ea e spe-
cies and phylogene ic di e si y han o he slope posi ions, highligh ing hei sig-
ni ican conse a ion alue o plan di e si y. Ka s dep essions, cha ac e ized
by deep, e ile soils, a e especially suscep ible o an h opogenic con e sion.
In sou hwes e n China, his has esul ed in subs an ial na u al o es loss and
deg ada ion, p ima ily due o hei con e sion in o ag icul u al and plan a ion
lands (Yuan e al. 2024). The e o e, conse a ion e o s o o es s in ka s
dep essions should be p io i ized. Ou indings p o ide c i ical insigh s o plan-
ning o es ese es and de eloping conse a ion s a egies in ka s egions.
Conclusion
The esul s show ha species di e si y in he he b and sh ub laye s ini ially de-
c eases and hen inc eases. In con as , species ichness in he ee laye ini ial-
ly inc eases and hen dec eases, while he emaining indices show an o e all
inc easing end in he TKSR along s eep ka s hillslopes. S epwise eg ession
analysis e ealed ha SA, SP, and SOM all ha e high explana o y powe o spe-
cies di e si y in he he b laye . BA, SA, and soil pH explained much o he a ia-
ion in species di e si y in he sh ub laye , whe eas BA, CD, and SP had s ong
in luences on species di e si y in he ee laye . Phylogene ic di e si y in he he b,
sh ub, and ee laye s ini ially dec eased and hen inc eased as he slope g adien
inc eased. Plan communi ies exhibi ed ela i ely high phylogene ic di e si y in
he dep essions and on he uppe slopes, sugges ing longe e olu iona y his o-
ies and g ea e e olu iona y di e si y and a ia ion. In con as , plan communi-
ies on he lowe and middle slopes had ela i ely lowe phylogene ic di e si y.
SA and SOM collec i ely had signi ican e ec s on he phylogene ic di e si y o
he he b laye ; SA, NO3
--N, and BA signi ican ly in luenced he phylogene ic di-
e si y o he sh ub laye ; and SP, ROR, BA, and CD had signi ican e ec s on he
phylogene ic di e si y o he ee laye . The he b and ee laye s exhibi ed phylo-
gene ic clus e ing, and habi a il e ing was he p ima y ecological p ocess in lu-
encing communi y assembly. The sh ub laye exhibi ed phylogene ic o e dispe -
sion, and compe i i e exclusion was he p ima y ecological p ocess in luencing
communi y assembly. BA, ROR, and soil pH we e he p ima y ac o s in luencing
90
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Qingling Pang e al.: Plan di e si y on s eep ka s hillslopes
he phylogene ic s uc u e o he he b laye ; soil pH, NH4
+-N, and CD s ongly in-
luenced he phylogene ic s uc u e o he sh ub laye ; and soil EMg, ECa, pH, and
SP had high explana o y powe o he phylogene ic s uc u e o he ee laye .
O e all, he esul s show ha de e minis ic p ocesses (habi a il e ing and com-
pe i i e exclusion) ha e signi ican impac s on communi y assembly in he TKSR;
howe e , some communi ies a e also in luenced by neu al p ocesses. Fu he -
mo e, dep essions and uppe slopes ha e highe conse a ion alue because o
hei highe le els o species and phylogene ic di e si y in sou hwes e n China.
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
This wo k was suppo ed by he Guangxi Na u al Science Founda ion (2021GXNSF-
FA196005, 2021GXNSFAA196024, 2022GXNSFBA035633, 2022GXNSFBA035461); he
Special Funding o Guangxi Bagui Young Top Talen s P og am ( o Zhonghua Zhang);
and he Na ional Na u al Science Founda ion o China (31960275, 31760128).
Au ho con ibu ions
Qingling Pang: Concep ualiza ion; da a cu a ion; me hodology; w i ing—o iginal d a . Cong
Hu: Funding acquisi ion; supe ision; w i ing—o iginal d a . Chaohao Xu: supe ision; w i -
ing—o iginal d a . Chao ang Zhong: supe ision; w i ing—o iginal d a . Zhonghua Zhang:
Concep ualiza ion; unding acquisi ion; supe ision; isualiza ion; w i ing— e iew and edi -
ing. Gang Hu: P ojec adminis a ion; supe ision; isualiza ion; w i ing— e iew and edi ing.
Au ho ORCIDs
Qingling Pang h ps://o cid.o g/0009-0002-1168-7402
Cong Hu h ps://o cid.o g/0000-0001-7507-4520
Chaohao Xu h ps://o cid.o g/0000-0002-0437-5390
Chao ang Zhong h ps://o cid.o g/0000-0002-3509-3628
Zhonghua Zhang h ps://o cid.o g/0000-0003-2094-698X
Gang Hu h ps://o cid.o g/0000-0002-0662-811X
Da a a ailabili y
All da a ha suppo he indings o his s udy a e a ailable upon eques .
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Supplemen a y ma e ial 1
En i onmen al a iables along he slope posi ion g adien in a ka s peak-
clus e dep ession land o m
Au ho s: Qingling Pang, Cong Hu, Chaohao Xu, Chao ang Zhong, Zhonghua Zhang, Gang Hu
Da a ype: docx
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