His o ical biogeog aphy and cha ac e -mapping
o Ac idoca pus (Malpighiaceae) e idence a e ised
in agene ic classi ica ion sys em and shi s in A ican
biomes om he Eocene o he Miocene
Ra ael F. de Almeida1,3, Sede a No o iana Rasambo2, Kelda Ellio 1, Ma co O.O. Pelleg ini1,3,
Ni ohenin soa Rako oni ina2, Tiana Rand iamboa onjy2, Mamy Tiana Rajaonah2,
Mian sa And ianan enaina2, No o Feni a Rand iana imanana2, Nan enaina He izo
Rako omalala2, Mbola Rako ond a simba2, Sede a Ny Aina Ranai oson2, Ta i aniaina
Rand ia sa azaka2, Da id Goyde 1, Ma ia S. Vo on so a1
1 Royal Bo anical Ga dens, Kew, Richmond, London, Su ey, UK
2 Kew Madagasca Conse a ion Cen e, An anana i o, Madagasca
3 C.E.Moss He ba ium, Uni e si y o he Wi wa e s and, Johannesbu g, Sou h A ica
Co esponding au ho : Ra ael F. de Almeida ([email p o ec ed])
Academic edi o : Isabel La idon ♦
Recei ed
13 May 2025 ♦
Accep ed
29 Augus 2025 ♦
Published
4 No embe 2025
Abs ac
Backg ound and aims – Ac idoca pus belongs o one o he se en Malpighiaceae lineages ha dispe sed om he
Neo opics o he Paleo opical egion, being by a he mos widely di e si ied and dis ibu ed genus o he amily in
A ica. In his s udy, we es ed he monophyly and alidi y o he cu en in agene ic classi ica ion o Ac idoca pus wi h
a da ed molecula phylogeny. We also econs uc ed ances al ange dis ibu ions o biomes and con inen s o elucida e
which ou e led o he colonisa ion o A ica by he mos ecen common ances o (MRCA) o his genus.
Ma e ial and me hods – We sampled six genes (ITS, PHYC, ma K, ndhF, bcL, and nL-F), 21 species o Ac idoca pus,
and h ee ou g oup species o es he monophyly o he in agene ic classi ica ion o he genus. BI and ML analyses
we e pe o med o he combined molecula da ase . A o al o 20 mo phological cha ac e s we e op imised on he ee.
Calib a ion poin s de i ed om a published Malpighiaceae ch onog am we e used o a da ing analysis. Ances al a eas
o Ac idoca pus and i s ela i es we e es ima ed o con inen al (Sou h Ame ica, A ica, India, Madagasca , and New
Caledonia) and biome (d y o es s, humid o es s, and sa annas) anges.
Key esul s – The p e-exis ing in agene ic classi ica ion o Ac idoca pus was eco e ed as non-monophyle ic due o
being solely based on homoplas ic mo phological cha ac e s. The MRCA o Ac idoca pus colonised ain o es s o Eas
A ica + Madagasca 43 Mya ia he Gondwana ou e and g ea ly di e si ied in his egion, wi h a single long-dis ance
dispe sal e en om Madagasca o New Caledonia (Oceania). The genus colonised A ican d y o es s a leas ou
di e en imes, s a ing in he Oligocene and di e si ied a single ime in Malagasy sa annas in he Miocene.
Keywo ds
ana omy, Ac idoca peae, Malpighiales, mo phology, Paleo opics, sys ema ics
INTRODUCTION
Ac idoca pus Guill., Pe . & A.Rich. (Ac idoca peae
R.F.Almeida) is he la ges genus o A ican Malpighiaceae,
comp ising 36 species o ees, sh ubs, o lianas (Almeida
e al. 2024a; POWO 2025). The genus is cu en ly di ided
in o wo subgene a (A. subg. Ac idoca pus and A. subg.
Anophylla is Nied.), ou sec ions (sec . Ac idoca pus,
Plan Ecology and E olu ion 158 (3): 428–444, 2025
h ps://doi.o g/10.5091/plece o.158824
Copy igh Ra ael F. de Almeida, Sede a No o iana Rasambo, Kelda Ellio , Ma co O.O. Pelleg ini, Ni ohenin soa Rako oni ina, Tiana Rand iamboa onjy,
Mamy Tiana Rajaonah, Mian sa And ianan enaina, No o Feni a Rand iana imanana, Nan enaina He izo Rako omalala, Mbola Rako ond a simba, Sede a Ny
Aina Ranai oson, Ta i aniaina Rand ia sa azaka, Da id Goyde , Ma ia S. Vo on so a. This is an open access a icle dis ibu ed unde he e ms o he C ea i e
Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed 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.
Plan Ecology and E olu ion is published by Meise Bo anic Ga den and he Royal Bo anical Socie y o Belgium.
RESEARCH ARTICLE
Plan Ecology and E olu ion 158 (3): 428–444, 2025 429
Diac a, Mac an he a, and Mic an he a), wo subsec ions
(subsec . Adenoch ea Nied. and Adenolepis Nied.), ou
se ies (se . Hemioop e ys Nied., Lep o hachis Nied.,
Machae op e ys Nied., and Pachy hachis Nied.), and i e
subse ies (subse . Ac inadenia Nied., Anadenia Nied.,
Monadenia Nied., Pleiadenia Nied., and Zygadenia Nied.;
Suppl. ma e ial 3; Niedenzu 1928). Niedenzu (1928) based
his in agene ic classi ica ion, sligh ly modi ied by A ènes
(1945) and Mo on (1968), on mo phological cha ac e s
such as habi ype, shape o in lo escence axis, p esence
and numbe o glands in b ac eoles and sepals, leng h o
an he s, pos u e o he do sal wing, and shape o he nu s
in me ica ps.
Species o Ac idoca pus a e commonly ound in
sa annas, seasonally d y o es s, and ain o es s om
I an o he A abian Peninsula, sub-Saha an A ica, and
Madagasca (Almeida e al. 2024a; POWO 2025). A single
species o his genus, A. aus ocaledonicus Baill., p esen s
a disjunc dis ibu ion on he island o New Caledonia
(Oceania; POWO 2025). This genus belongs o one o
he se en lineages o Malpighiaceae ha dispe sed and
di e si ied in he Paleo opical egion om mid-Eocene
o la e Pliocene (Da is e al. 2014). P e ious his o ical
biogeog aphy s udies o his genus e ealed ha he i s -
di e ging lineage comp ises only species endemic o
Madagasca and New Caledonia, while he species om
he emaining lineages a e g ouped in o a la ge clade
wi h species con ined o con inen al A ica (Da is e
al. 2002; Da is and Ande son 2010). These s udies also
indica e ha he o igin o Ac idoca pus di e si ica ion
is in Sou heas e n A ica, wi h successi e mig a ions o
Wes A ica and hen back o No heas e n A ica (Da is
e al. 2002).
The i s s udy o es he monophyly o he genus
is Da is e al. (2002), which used he ITS, ndhF, and
nL-F ma ke s. Ac idoca pus was eco e ed as a sis e
g oup o he Asian endemic genus B achylophon Oli .
The Ac idoca pus + B achylophon clade has consis en ly
been eco e ed as he second lineage o di e ge in
Malpighiaceae (Came on e al. 2001; Da is e al. 2001;
Da is and Ande son 2010; Almeida e al. 2024a).
Following hese, a molecula gene ic phylogeny o
Malpighiaceae based on PHYC, ma K, ndhF, and bcL
ma ke s was published (Da is and Ande son 2010).
I eco e ed he same opology as Da is e al. (2002)
bu wi h ewe species sampled. Bo h s udies sampled
a ound 16 accep ed species o Ac idoca pus, including i s
ype species, A. plagiop e us Guill. & Pe . E en hough
bo h s udies sampled species om wo subgene a and
di e en sec ions and subsec ions cu en ly accep ed in
Ac idoca pus, no discussion was e e p esen ed ega ding
he monophyly o he in agene ic anks wi hin he genus.
In he cu en s udy, we econs uc he molecula
phylogeny o Ac idoca pus o assess he monophyly o he
p e-exis ing in agene ic anks wi hin he genus, based
on ITS, PHYC, ma K, ndhF, bcL, and nL-F ma ke s. A
single species o B achylophon was sampled as he sis e
g oup, and By sonima Rich. ex Kun h and Galphimia Ca .
as ou g oups o he analyses. The esul ing ee was hen
used o op imise 20 mac o- and mic omo phological
cha ac e s, which we e es ed o homoplasies and
synapomo phies. Addi ionally, we da ed he molecula
phylogeny o econs uc he ances al anges o
Ac idoca pus a con inen al and biome le els, he eby
elucida ing i s biogeog aphic his o y in A ica o e ime.
Mo e speci ically, we aimed o answe he ollowing
ques ions: Which con inen al ou e led he Mos Recen
Common Ances o (MRCA) o Ac idoca pus o colonise
con inen al A ica and Madagasca ? In con inen al A ica
and Madagasca , which biomes did he MRCA o he
genus di e si y in?
MATERIAL AND METHODS
Molecula analyses
We included 21 species o Ac idoca pus (58% o he 36
accep ed species), each ep esen ed by a single specimen,
sampling all accep ed subgene a and sec ions. A single
species o B achylophon was included as he sis e g oup,
and By sonima and Galphimia as ou g oups o ou
analyses, acco ding o Da is and Ande son (2010) and
Almeida e al. (2024a). All sequences o he ITS, PHYC,
ma K, ndhF, bcL, and nL-F ma ke s we e e ie ed
om GenBank (Suppl. ma e ial 1), edi ed using Geneious
.4.8.4 (Kea se e al. 2012), and aligned using Muscle
.1.0 (Edga 2004), wi h subsequen adjus men s in he
p elimina y ma ices made manually by eye.
Phylogene ic analyses we e pe o med using Bayesian
in e ence and Maximum Likelihood. The model selec ion
used hie a chical likelihood a io es s (HLRT) on
jModel es 2 (Da iba e al. 2012). Bo h model-based
me hods we e ca ied ou wi h mixed models (GTR+G+I)
and unlinked pa ame e s, using M Bayes .3.1.2 (Ronquis
and Huelsenbeck 2003) and RAxML .8 (S ama akis 2014)
and implemen ed on axmlGUI .2 (Edle e al. 2021).
Fo he Bayesian in e ence, he Ma ko Chain Mon e
Ca lo (MCMC) was ca ied ou in wo simul aneous
independen uns wi h ou chains each (one cold and
h ee hea ed), sa ing one ee e e y 1,000 gene a ions o
a o al o en million gene a ions. We excluded 20% o
he e ained ees as bu n-in and checked o a s a iona y
phase o likelihood and he E ec i e Sample Size (ESS)
alues highe han 200 o all pa ame e s on T ace .1.7
(Rambau e al. 2018). The pos e io p obabili ies (PP) o
clades we e based on he majo i y- ule consensus, using
he s o ed ees, and calcula ed wi h M Bayes .3.1.2
(Ronquis and Huelsenbeck 2003).
Mo phological analyses
Fo all species in he s udy, en mac omo phological
cha ac e s we e sco ed om Niedenzu (1928), A ènes
(1945, 1954), and Almeida e al. (2020), and en
mic omo phological cha ac e s we e sco ed om
Guesdon e al. (2019) and San os e al. (2023). Cha ac e
Almeida e al.: His o ical biogeog aphy and cha ac e -mapping o Ac idoca pus (Malpighiaceae)430
coding ollowed he ecommenda ions o Se eno
(2007) o mo phological analyses. P ima y homology
hypo heses (De Pinna 1991) we e p oposed o 20 mac o-
and mic omo phological cha ac e s (Suppl. ma e ial 2).
All cha ac e s we e op imised on he conca ena ed ee
using he Maximum Likelihood unc ion (mk1 model)
o Mesqui e .3.61 (Maddison and Maddison 2019) and
isualised on Winclada (Nixon 1999).
Calib a ion
Da ing es ima es we e based on a simpli ied ul ame ic
Bayesian combined ee gene a ed wi h BEAST .1.8.4
(D ummond e al. 2012). This analysis used a elaxed,
unco ela ed logno mal clock and Yule p ocess specia ion
p io o in e ing ees. The calib a ion pa ame e s
we e based on p e ious es ima es de i ed om a
comp ehensi e s udy o he whole Malpighiaceae (Da is
e al. 2014) because no known ossil o Ac idoca pus was
a ailable in he li e a u e. We op ed o calib a ing a he
c own node o Ac idoca pus using a no mal p io wi h
mean ini ial alues o 43.0 Mya and a s anda d de ia ion
o 1.0 (Da is e al. 2014). Two sepa a e and con e gen
uns we e ca ied ou , wi h 10,000,000 gene a ions,
sampling e e y 1,000 s eps and 2,000 ees as bu n-in. We
checked o ESS alues highe han 400 o all pa ame e s
on T ace .1.7 (Rambau e al. 2018). T ee opology was
assessed by selec ing he maximum clade c edibili y ee
o he uns wi h T ee Anno a o and he g aphic display
gene a ed in FigT ee .1.4.4 (Rambau 2018).
Ances al a ea econs uc ion
Species dis ibu ion da a we e compiled om POWO
(2025) and GBIF (2025). Ances al a eas o Ac idoca pus
and i s ela i es we e es ima ed o con inen al (Sou h
Ame ica, A ica, India, Madagasca , and New Caledonia)
and biome (d y o es s, humid o es s, and sa annas)
anges using he R package (R Co e Team 2025)
BioGeoBEARS .1.1.1 (Ma zke 2018) and he models
DEC, DIVALIKE, and BAYAREALIKE. Conside ing he
possibili y o unde es ima ion o ex inc ion due o he
null anges o species allowed by he models, analyses
wi h he “*” op ion we e also pe o med. The “j” op ion
was used o conside ounde specia ion e en s. Di e en
combina ions o he h ee models and he wo op ions
gene a ed a o al o 12 models. The bes - i ing models o
analyses we e de e mined by AIC weigh a ios.
RESULTS
Phylogene ic analyses
The combined da ase included 6,110 nucleo ides. The e
we e no majo incong uences in he opologies p oduced
by BI and ML analyses, so we chose o p esen he esul s
on a single summa ised ee (Fig. 1). Ac idoca pus was
esol ed as monophyle ic and sis e o B achylophon,
wi h he By sonima + Galphimia clade as he ou g oup o
he analysis (Fig. 1). The BI analysis eco e ed an almos
en i ely esol ed ee, while he ML analysis eco e ed a
ully esol ed ee wi h mos clades well-suppo ed (> PP
95% / BS 70) and a ew poo ly suppo ed (< PP 95% / BS
70) (Fig. 1).
Wi hin Ac idoca pus, h ee well-suppo ed clades
we e eco e ed: A. subg. Anophylla is 1 (highligh ed
in pink on Fig. 1), A. subg. Ac idoca pus, and A. subg.
Anophylla is 2 (highligh ed in blue on Fig. 1). Only A.
subg. Ac idoca pus was eco e ed as monophyle ic, wi h
A. subg. Anophylla is eco e ed in o wo dis an clades
ep esen ing sec ions Mac an ha and Mic an ha (Fig. 1).
Fo bo h sec ions o Ac idoca pus, all subsec ions we e
eco e ed as pa aphyle ic (black and whi e squa es in Fig.
1).
Cha ac e mapping
Ac idoca pus subg. Anophylla is 1 (in pink on Fig.
1; = A. subg. Madagasca iensis) was eco e ed wi h
h ee synapomo phies ( ees (0/0), lea es abaxial side
uni o mly and densely hai y wi h epide mis hidden (1/1),
and cincinni b ac eoles 2-glandula (18/2)) and a single
homoplasy (lea es wi h mid ib ascula sys em in an open
a ch wi hou con olu ed ex emi ies (8/2)). Ac idoca pus
subg. Ac idoca pus (in whi e on Fig. 1) was eco e ed
wi h a single synapomo phy (lianas (0/2)) and homoplasy
(lea blade glands p ominen in ela ion o he epide mis
(17/2)). The clade comp ising A. subg. Ac idoca pus (in
whi e on Fig. 1) + A. subg. Anophylla is 2 (in blue on Fig.
1) was eco e ed wi h a single synapomo phy (sepals wi h
3 glands (19/3)). Finally, A. subg. Anophylla is 2 (in blue
on Fig. 1) was eco e ed wi h a single synapomo phy
(lea es wi h mid ib ascula sys em con o ma ion in
an open a ch wi h con olu ed ex emi ies (8/3)) and
homoplasy (lea epide mis wi h cuboid cells (2/0)).
Di e gence imes es ima ion and ances al ange
econs uc ions
BioGeoBEARS selec ed he DIVALIKE + J model o bo h
con inen al (LnL = -16.85) and biome (LnL = -23.56)
analyses due o he lowes LnL sco es (Figs 2, 3). The
MRCA o he B achylophon + Ac idoca pus clade (i.e.
ibe Ac idoca peae) a ose a 43 Mya and was widesp ead
in ain o es s h ough A ica, he Indian subcon inen ,
and Madagasca (Figs 2, 3). A he same ime, he MRCA
o B achylophon colonised he ain o es s o he Indian
subcon inen (Figs 2, 3). The MRCA o Ac idoca pus
a ose a 41 Mya and was widesp ead in ain o es s o
con inen al A ica and Madagasca (Figs 2, 3). A he same
ime, he MRCA o A. subg. Anophylla is 1 colonised he
ain o es s o Madagasca , di e si ying a 32 Mya (Figs 2,
3). The MRCA o A. humbe ii colonised he d y o es s
o Madagasca a he same age, while he MRCA o A.
adenopho us di e si ied in he ain o es s o Madagasca
a 22 Mya (Figs 2, 3). The MRCA o A. excelsus + A.
pe ie i + A. aus ocaledonicus clade di e si ied in he
Plan Ecology and E olu ion 158 (3): 428–444, 2025 431
Figu e 1. Phylogene ic ela ionships o Ac idoca pus based on ML and Bayesian analyses o ITS, PHYC, ma K, ndhF, bcL, and
nL-F sequences. B anch leng hs a e shown on he le . Cha ac e -mapping esul s a e shown on he igh , wi h ed numbe s on
he b anches ep esen ing PP alues (abo e) and boo s ap alues (below). Ci cles show homoplas ic cha ac e s (open ci cles)
and apomo phic cha ac e s ( ed ci cles). Pink clade = A. subg. Anophylla is 1 (= A. subg. Anophylla is sec . Mic an he a), black
squa es = A. subg. Anophylla is sec . Mic an he a subsec . Madagasca iensis, whi e squa es = A. subg. Anophylla is sec . Mic an he a
subsec . Neocaledonici. Blue clade = A. subg. Anophylla is 2 (= A. subg. Anophylla is sec . Anomalop e ys), black squa es = A. subg.
Anophylla is sec . Anomalop e ys subsec . Adenoch ea, whi e squa es = A. subg. Anophylla is sec . Anomalop e ys subsec . Adenolepis.
sa annahs o Madagasca om 17–10 Mya (Figs 2, 3). The
MRCA o A. aus ocaledonicus colonised New Caledonia
(Oceania) om a Malagasy ances o a 10 Mya (Figs 2,
3). The MRCA o Wes A ican Ac idoca pus (i.e. A.
subg. Ac idoca pus + A. subg. Anophylla is 2) colonised
con inen al A ican ain o es s a 39 Mya (Figs 2, 3).
Finally, he colonisa ion o con inen al A ican d y o es s
ook place a h ee di e en imes, s a ing a 39 Mya in
Almeida e al.: His o ical biogeog aphy and cha ac e -mapping o Ac idoca pus (Malpighiaceae)432
Figu e 2. Ch onog am and DIVALIKE + J ances al ange econs uc ions o con inen s in Ac idoca pus. Le : ec onic econs uc ion
om 40 Mya e idencing he possible dispe sal ou es o he MRCA o Ac idoca pus in he Ame ican con inen (da k blue, A),
Indian subcon inen (ligh blue, B), con inen al A ica (g een, C), Madagasca (yellow, D), and New Caledonia ( ed, E). Pho os: I.
B achylophon anas omosans C aib. by Phuhu T ang-Racha; II. Ac idoca pus excelsus by Cha les Rako o ao; III. Ac idoca pus s aud ii
by Ehoa n Bidaul ; IV. Ac idoca pus longi olius by Ehoa n Bidaul ; V. Ac idoca pus na ali ius by G aham G ie e; VI. Ac idoca pus
o ien alis by Da id Hacke Fische .
he MRCA o A. plagiop e us in Wes A ica, 24 Mya in
he MRCA o Ac idoca pus in Eas A ica, and 11 Mya in
he MRCA o he Wes A ican A. spec abilis (Figs 2, 3).
DISCUSSION
The molecula phylogeny o Ac idoca pus p esen ed
he ein di e s in species sampling and molecula ma ke s
om he s udies o Da is e al. (2002) and Da is and
Ande son (2010). We sampled all p e ious species and all
six genes ( wo nuclea and ou plas id) used by p e ious
s udies, as well as sequences o h ee addi ional species
(A. alopecu us, A. chlo op e us, and A. soco anus).
Fu he mo e, we only included a single specimen o each
accep ed species, signi ican ly educing he phylogene ic
noise in he Bayesian and Maximum likelihood analyses.
This had o be done o minimise he amoun o missing
da a be ween he ma ices eco e ed o he six ma ke s, a
s anda d phylogene ic me hod applied o seconda y da a.
In agene ic classi ica ion
The in agene ic classi ica ion o Ac idoca pus was
p oposed by Niedenzu (1928), who di ided he genus
in o wo subgene a (A. subg. Ac idoca pus and A.
subg. Anophylla is), ou sec ions (sec . Ac idoca pus,
Diac a, Mac an he a, and Mic an he a), wo subsec ions
(subsec . Adenoch ea Nied. and Adenolepis Nied.), ou
se ies (se . Hemioop e ys Nied., Lep o hachis Nied.,
Machae op e ys Nied., and Pachy hachis Nied.), and i e
subse ies (subse . Ac inadenia Nied., Anadenia Nied.,
Monadenia Nied., Pleiadenia Nied., and Zygadenia Nied.;
Suppl. ma e ial 3). This au ho based his in agene ic
classi ica ion on mo phological cha ac e s such as habi
ype, shape o in lo escence axis, p esence and numbe o
Plan Ecology and E olu ion 158 (3): 428–444, 2025 433
Figu e 3. Ch onog am and DIVALIKE + J ances al ange econs uc ions o biomes (colou ed map on he le ) in Ac idoca pus.
Black and g ey maps on he igh ep esen he o de o main colonisa ion e en s in A ica by Ac idoca pus. The highligh ed a ea in
he ch onog am ep esen s he Miocene age o Malagasy sa annas.
glands in b ac eoles and sepals, leng h o an he s, pos u e
o he do sal wing, and shape o he nu s in me ica ps.
Ac idoca pus. subg. Anophylla is comp ises ees o
sh ubs, wi h e ec b ac eoles, sho an he s, me ica ps
wi h an e ec do sal wing, and me ica p nu s semi-
len i o m in shape, while A. subg. Ac idoca pus comp ises
lianas wi h pa en b ac eoles, long an he s, me ica ps wi h
a pa en do sal wing, and me ica p nu s semi-globose o
semi-o oid in shape (Niedenzu 1928; Suppl. ma e ial
3). Ac idoca pus subg. Ac idoca pus is di ided in o wo
sec ions based on he shape and con luence o he apex
o an he s: A. subg. Ac idoca pus sec . Ac idoca pus
(Monac a) is cha ac e ised by he acu e apices o an he s
and conni en an he sacs, while A. subg. Ac idoca pus
sec . Diac a Nied. is comp ised o species wi h bico na e
apices o an he s and di e gen an he sacs (Niedenzu
1928; Suppl. ma e ial 3). Ac idoca pus subg. Anophylla is
is di ided in o wo sec ions based on he shape and
p esence o glands in he sepals and he leng h o an he s
and ilamen s: A. subg. Anophylla is sec . Mac an ha
Nied. wi h o a e- o und, 1–8 glandula sepals and
3.5–7 mm long an he s; and A. subg. Anophylla is sec .
Mic an ha Nied. wi h o a e, eglandula sepals and 1.5–2
mm long an he s (Niedenzu 1928). Niedenzu (1928)
di ided sec . Mac an ha in o wo subsec ions (subsec .
Adenoch ea Nied. and Adenolepis Nied.), ou se ies (se .
Hemioop e ys Nied., Lep o hachis Nied., Machae op e ys
Nied., and Pachy hachis Nied.) and i e subse ies (subse .
Ac inadenia Nied., Anadenia Nied., Monadenia Nied.,
Pleiadenia Nied., and Zygadenia Nied.; Suppl. ma e ial 3).
In con as , A ènes (1945) di ided sec . Mic an ha Nied.
in o wo subsec ions (subsec . Madagasca iensis and
Neocaledonici A ènes; Suppl. ma e ial 3).
All diagnoses p esen ed o Niedenzu’s p oposed
subgene a we e no compa able o each o he , making i
inc edibly di icul o co ec ly assign any newly published
species o his in agene ic ank ci cumsc ip ions. These
incompa able diagnoses we e also poin ed ou by A ènes
(1945), who published wo addi ional subsec ions
o A. subg. Anophylla is sec . Mic an ha. Subsec ion
Madagasca iensis A ènes included all Ac idoca pus
species om Madagasca , while subsec . Neocaledonici
A ènes only included he New Caledonia endemic
A. aus ocaledonicus. A ènes based his in agene ic
Almeida e al.: His o ical biogeog aphy and cha ac e -mapping o Ac idoca pus (Malpighiaceae)434
classi ica ion on mo phological cha ac e s such as habi
ype, p esence, shape, and size o glands in b ac eoles,
pe al consis ency and shape, and leng h o ilamen s
and an he s (A ènes 1945). La e , Mo on (1968) ound
ou ha A. subg. Anophylla is sec . Mac an he a was an
illegi ima e name due o pa o i s species being p e iously
published unde Anomalop e ys (DC.) G.Don. Thus,
he published A. subg. Anophylla is sec . Anomalop e ys
o accommoda e his name and synonymised A. subg.
Anophylla is sec . Mac an he a unde i .
Ou esul s showed ha om all in agene ic anks
cu en ly accep ed in Ac idoca pus, only A. subg.
Ac idoca pus, A. subg. Anophylla is sec . Anomalop e ys,
and A. subg. Anophylla is sec . Mic an ha we e esol ed as
monophyle ic (Fig. 1). All emaining in agene ic anks
we e ound o lack monophyly. Mos o he diagnos ic
mo phological cha ac e s ha we e used o hei
ci cumsc ip ion we e eco e ed as highly homoplas ic
(Fig. 1). Ou analyses ha e eco e ed he p esence o
nec a -sec e ing sepal glands as a synapomo phy o
con inen al A ican Ac idoca pus (Fig. 1). None heless,
mos Paleo opical lineages o Malpighiaceae also p esen
nec a -sec e ing sepal glands, esul ing om a pollina ion
shi ha occu ed in he ances o o hese lineages when
dispe sing om he Neo opics o he Paleo opics (Da is
e al. 2014). Sepal glands in species o Ac idoca pus
a e small and es ic ed o he base nea ma gins o
a e a single, la ge, and in e sepaline-placed gland
esul ing om he usion o glands om adjacen sepals
(Guesdon e al. 2019). Thus, nec a -sec e ing glands a e
a homoplas ic cha ac e in Malpighiaceae ela ed o a
pollina ion shi and canno be ega ded as an exclusi e
ai o Ac idoca pus.
A eci cumsc ip ion o he in agene ic classi ica ion
o Ac idoca pus is p esen ed in he axonomy sec ion
below.
His o ical biogeog aphy
P e ious s udies (Da is e al. 2002) jus using dispe sal-
ica iance analysis poin ed o he bo eo opical ou e as
he dispe sal ou e aken by he MRCA o Ac idoca pus.
Howe e , ou esul s poin o a Gondwana dispe sal ou e,
suppo ed by 12 di e en s a is ical models (Ma zke
2018) and a new ossil eco d (Ali e al. in p ess). Ou
esul s indica e ha he MRCA o Ac idoca pus colonised
ain o es s o A ica and Madagasca ia he Gondwana
ou e. This is no su p ising since 35 species (ou o 36
species) o he genus a e cu en ly es ic ed o coun ies
in con inen al A ica and Madagasca wi h pa ches o
ain o es (POWO 2025). Ou analyses also co obo a e
he exis ence o a long-dis ance dispe sal e en a 43 Mya
o he MRCA o Ac idoca pus + B achylophon om i s
Neo opical ela i es (i.e. sub amily By sonimoideae),
co obo a ing Da is e al. (2014). This dispe sal e en
occu ed om he Ame icas o he Indian subcon inen
+ Madagasca + A ica h ough he Gondwana ou e
ins ead o he Bo eo opical ou e (Da is e al. 2004,
2014). The Bo eo opical ou e is g ea ly suppo ed by
ossils (Gi nish and Renne 2004) and he calib a ed
phylogenies o se e al g oups o plan s, such as Clusiaceae
(Mesegue e al. 2018), Annonaceae (Thomas e al. 2015),
Symplocaceae (F i sch e al. 2015), Leguminosae (Es ella
e al. 2017), Lycopodiaceae (Bau e e al. 2018), and
Malpighiaceae (Da is e al. 2014; Almeida and an den
Be g 2022), among o he s.
The MRCA o Ac idoca pus, ep esen ed by he
Ac idoca pus c own node, s a ed o di e si y in
ain o es s o he A ican con inen a ound 43 Mya, wi h
a leas ou di e en colonisa ion e en s in d y o es s
and a single colonisa ion e en in sa annas. These esul s
co obo a e he p esence o a pan-A ican ain o es om
Wes o Eas du ing he Eocene (Cou eu e al. 2021),
which was likely he mig a ion ou e o he MRCA
o con inen al A ican Ac idoca pus o dispe se om
Madagasca o Wes A ica and back again o con inen al
Eas A ica (Fig. 3). This is a well-documen ed pa e n in
A ican ain o es s, which subs an ia es he b eakup o
his con inuous pan-A ican ain o es in he coole and
d ie Miocene (Cou eu e al. 2021). Ou esul s exhibi
ha mos d y o es lineages in Ac idoca pus s a ed o
di e si y in he Oligocene, ma ching he Oligocene o igins
o mos neo opical d y o es lineages o Malpighiaceae,
bo h in Cen al and Sou h Ame ica (Willis e al. 2014;
Almeida e al. 2018). The same Eocene/Oligocene pa e n
o A ican d y o es s di e si ica ion is also obse ed in
Apocynaceae (Bi encou e al. 2021), Zygophyllaceae (Wu
e al. 2018), and Rhamnaceae (Rickenback e al. 2024).
Addi ionally, he only sa anna lineage o Ac idoca pus
di e si ied h oughou he en i e Miocene in Madagasca ,
co obo a ing he Miocene o igins o Ame ican, A ican,
and Malagasy sa anna species o Malpighiaceae (Almeida
e al. 2018, 2024b; Almeida and an den Be g 2020). The
ema kable cong uence o New and Old Wo ld sa anna
o igins has al eady been s a ed by Penning on and
Hughes (2014), bu addi ional da ed phylogenies a e s ill
needed o p ope ly co obo a e he imeline o he o igins
o sa annas wo ldwide.
Nume ous s udies on he age o Malagasy sa annas
can be ound in he li e a u e, d awing on a ious ypes
o e idence. Based on da ed molecula phylogenies o C3
g asses, some au ho s showed ha sa annas o igina ed
in he Oligocene o Miocene (Hackel e al. 2018). The
C3 g asses con inued o adia e in o he Pliocene and
Qua e na y, while C4 g asses o igina ed la e in he
Miocene o Pliocene (8–3 Mya) and adia ed expansi ely
in o he Pleis ocene (Hackel e al. 2018; Hagl e al. 2021).
The Malagasy g ass lineages a e linked o dispe sal e en s
p ima ily om mainland A ica, a pa e n also obse ed
in Ac idoca pus. Mo eo e , he adia ion o C4 g asses in
Madagasca coincided wi h he o igin and global sp ead
o open, g assy, i e-p one landscapes du ing he Miocene
(Ce ling e al. 1997; Edwa ds e al. 2010; S ömbe g 2011).
Silande J e al. (2023) e idenced 41 woody plan lineages
endemic o he g assy ecosys ems o Madagasca . F om
his ini ial lis , we we e able o ind s udies wi h da ed
Plan Ecology and E olu ion 158 (3): 428–444, 2025 435
molecula phylogenies o a leas 14 na i e lineages
(A ecaceae, Aspa agaceae, As e aceae, Cype aceae,
Fabaceae, Malpighiaceae, Melas oma aceae, Oleaceae,
and Poaceae) showing a dis inc pa e n o Miocene
adia ion in Malagasy sa annas (Table 1).
TAXONOMIC TREATMENT
Ac idoca pus Guill. & Pe ., nom. cons. (Guillemin and
Pe o e 1831: 123)
Anomalop e ys (DC.) G.Don (Don 1831: 647) –
He e op e ys sec . Anomalop e ys DC. (De Candolle
1824: 592) – Type species: Anomalop e ys spica a
G.Don, nom. supe l. [Ac idoca pus smea hmannii
(DC.) Guill. & Pe .].
Rhinop e yx Nied. (Niedenzu 1896: 352) – Type species:
Rhinop e yx spec abilis Nied. [Ac idoca pus spec abilis
(Nied.) Doo n-Hoekm.].
Type species. Ac idoca pus plagiop e us Guill. & Pe .
Diagnosis. Ac idoca pus is cha ac e ised by i s ee, sh ub
o lianescen habi , lea es al e na e, usually abaxially
glandula nea he mid ein and base, long o sho
hy ses, b ac eoles 0–2 glandula , sepals 0–5 glandula ,
nec a -sec e ing, pos e io pe als 2, e ec , la e al pe als
pa en , all pe als wi h ma gins c ena e o den a e, s amens
10, ilamen s he e omo phic in leng h, an he s po icidal,
pollen pa asyncolpo a e, gynoecium 3-ca pella e, s yles
2, e ec and cu ed inwa ds, me ica ps wi h do sal wing
well-de eloped, and la e al wings absen (Niedenzu 1928).
Dis ibu ion. Ac idoca pus is cu en ly composed o 36
accep ed species (19 o which a e h ea ened; POWO
2025) dis ibu ed in con inen al A ica, Madagasca ,
he A abian Peninsula, I an, and Oceania (i.e. New
Caledonia) (Almeida e al. 2024a; POWO 2025).
Habi a and ecology. Ac idoca pus comp ises species o
ees, sh ubs, scanden sh ubs, o lianas ha a e endemic
o ain o es s, sa annas, and seasonally d y opical
o es s (Almeida e al. 2024a; POWO 2025).
No es. The e is no upda ed iden i ica ion key o all
species o Ac idoca pus, bu Niedenzu’s (1928) ea men
co e s 25 ou o he 36 cu en ly accep ed species.
Key o he subgene a o Ac idoca pus
1. T ees, lea blades abaxially, uni o mly, and densely hai y, epide mis hidden, ascula sys em in an open a ch wi hou con olu ed
ex emi ies in he mid ib po ion, b ac eoles 2-glandula , sepals eglandula , an he s 1–2 mm long, Madagasca , New Caledonia ...
.................................................................................................................................................................................... A. subg. Madagasca iensis
– Sh ubs, scanden sh ubs o lianas, lea blades glab ous o glab escen , epide mis isible, mid ib con ou plane-con ex o conca e-
con ex, b ac eoles 0–1-glandula , sepals 1-many-glandula , an he s 3.5–7 mm long, con inen al A ica, A abian Peninsula, I an ..
...................................................................................................................................................................................................................................2
2. Lianas, lea glands p ominen , epide mal cells elonga ed, ascula sys em bicon ex o plane-con ex in he mid ib po ion, b ac
and b ac eoles pa en , la e al pe als cuculla e, ma gins e ose, ilamen s sho e han an he , me ica ps wi h a pa en do sal wing,
me ica p nu s semi-globose o semi-o oid in shape ..................................................................................................A. subg. Ac idoca pus
– Sh ubs o scanden sh ubs, lea glands imp essed, epide mal cells cuboid, ascula sys em in an open a ch wi h con olu ed
ex emi ies in he mid ib po ion, b ac and b ac eoles e ec , la e al pe als plane, ma gins den a e o c ena e, ilamen s longe han
an he s, me ica ps wi h an e ec do sal wing, me ica p nu s semi-len i o m in shape ........................................A. subg. Anophylla is
Table 1. Compila ion o da ed molecula phylogenies a ailable in he li e a u e o he g oups o woody plan s na i e o Malagasy
woodland and g asslands, acco ding o Silande J e al. (2023).
Taxa Mya Re e ences
Malagasy Cype aceae 1 20 La idon e al. (2021)
Bo assus madagasca iensis (Jum. & H.Pe ie ) Boje ex Jum. & H.Pe ie (A ecaceae) 18 Bello e al. (2020)
T ibe Sone ileae (Melas oma aceae) 16 Ve anso-Libalah e al. (2018)
B andzeia ilici olia Baill. (Fabaceae) 15 Choo e al. (2020)
Memecylon buxi olium Blume (Melas oma aceae) 15 Ama asinghe e al. (2021)
Malagasy Poaceae 15 Hackel e al. (2018)
Dis ephanus Cass. (As e aceae) 14 Gos el e al. (2024)
Rousseauxia DC. (Melas oma aceae) 13 Ve anso-Libalah e al. (2018)
Ampho ocalyx Bake (Melas oma aceae) 13 Ve anso-Libalah e al. (2018)
Dionycha Naudin (Melas oma aceae) 13 Ve anso-Libalah e al. (2018)
Philgamia Baill. + Sphedamnoca pus Planch. ex Ben h. & Hook. . (Malpighiaceae) 13 Da is e al. (2014)
Sub ibe Ledebou iinae (Aspa agaceae) 10 Howa d e al. (2023)
Malagasy Cype aceae 2 10 La idon e al. (2021)
No onhia low yi Hong-Wa (Oleaceae) 7 Salmona e al. (2020)
Almeida e al.: His o ical biogeog aphy and cha ac e -mapping o Ac idoca pus (Malpighiaceae)436
1. Ac idoca pus subg. Ac idoca pus (Niedenzu 1921:
17)
Fig. 4
Ac idoca pus subg. Ac idoca pus sec . Ac idoca pus
(Niedenzu 1921: 17)
Ac idoca pus subg. Ac idoca pus sec . Diac a Nied.
(Niedenzu 1921: 17)
Type species. Ac idoca pus plagiop e us Guill. & Pe .
Diagnosis. Ac idoca pus subg. Ac idoca pus is
cha ac e ised by i s liana habi , lea blades glab ous o
glab escen , epide mis isible, lea glands p ominen ,
epide mal cells elonga ed, ascula sys em bicon ex o
plano-con ex in he mid ib po ion, b ac and b ac eoles
pa en , b ac eoles 0–1-glandula , sepals 1–many-
glandula , la e al pe als cuculla e, ma gins e ose, ilamen s
sho e han an he , me ica ps wi h a pa en do sal wing,
me ica p nu s semi-globose o semi-o oid in shape.
Dis ibu ion. Ac idoca pus subg. Ac idoca pus comp ises
six accep ed species dis ibu ed in Cen al and Wes e n
con inen al A ica.
Habi a and ecology. Species o Ac idoca pus subg.
Ac idoca pus a e ound in d y o ain o es s.
1.1. Ac idoca pus came unensis Nied. (Niedenzu 1915:
52)
1.2. Ac idoca pus ka angensis De Wild. (De Wildeman
1902: 27)
Ac idoca pus u escens Hu ch. (Hu chinson 1946: 520)
1.3. Ac idoca pus mac ocalyx Engl. (Engle 1905: 250)
1.4. Ac idoca pus mayumbensis Gonç. & E.Laune
(Gonçal es and Laune 1984: 141)
1.5. Ac idoca pus plagiop e us Guill. & Pe . (Guillemin
and Pe o e 1831: 123)
Ac idoca pus hi undo S.Moo e (Moo e 1880: 1)
Anomalop e ys obo a a G.Don (Don 1831: 647)
1.6. Ac idoca pus s aud ii (Engl.) Engl. ex Hu ch. &
Dalziel (Hu chinson and Dalziel 1928: 271)
Ac idoca pus smea hmannii a . s aud ii Engl. (Engle
1905: 251)
2. Ac idoca pus subg. Anophylla is Nied. (Niedenzu
1921: 3)
Fig. 5
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
(DC.) Mo on (Mo on 1968: 315)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenoch ea Nied. (Niedenzu 1921: 4)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenoch ea se . Hemioop e ys Nied. (Niedenzu
1921: 4)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenoch ea se . Hemioop e ys subse .
Ac inadenia Nied. (Niedenzu 1921: 4)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenoch ea se . Hemioop e ys subse .
Zygadenia Nied. (Niedenzu 1921: 5)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenoch ea se . Machae op e ys Nied.
(Niedenzu 1921: 7)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenoch ea se . Machae op e ys subse .
Anadenia Nied. (Niedenzu 1921: 7)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenoch ea se . Machae op e ys subse .
Monadenia Nied. (Niedenzu 1921: 13)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenoch ea se . Machae op e ys subse .
Pleiadenia Nied. (Niedenzu 1921: 8)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenolepis Nied. (Niedenzu 1921: 14)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenolepis se . Lep o hachis Nied. (Niedenzu
1921: 15)
Ac idoca pus subg. Anophylla is sec . Anomalop e ys
subsec . Adenolepis se . Pachy hachis Nied. (Niedenzu
1921: 14)
Ac idoca pus subg. Anophylla is sec . Mac an he a Nied.
(Niedenzu 1921: 4), nom. illeg.
He e op e ys sec . Anomalop e is DC. (De Candolle 1824:
592), nom. ej.
Type species. Ac idoca pus na ali ius A.Juss.
Diagnosis. Ac idoca pus subg. Anophylla is is
cha ac e ised by i s sh ub o scanden sh ub habi , lea
blades glab ous o glab escen , epide mis isible, lea
glands imp essed, epide mal cells cuboid, ascula sys em
in an open a ch wi h con olu ed ex emi ies in he mid ib
po ion, b ac and b ac eoles e ec , 0–1-glandula , sepals
1-many-glandula , la e al pe als plane, ma gins den a e o
c ena e, ilamen s longe han an he s, me ica ps wi h an
e ec do sal wing, me ica p nu s semi-len i o m in shape.
Dis ibu ion. Ac idoca pus subg. Anophylla is comp ises
22 accep ed species dis ibu ed in Eas A ica, he A abic
Peninsula, and I an.
Habi a and ecology. Species o Ac idoca pus subg.
Anophylla is a e ound in d y o es s o ain o es s.
2.1. Ac idoca pus alopecu us Sp ague (Sp ague 1909a:
185)
Ac idoca pus alopecu us a . alopecu us Sp ague (Sp ague
1909a: 185)
Ac idoca pus alopecu us a . machae op e us Nied.
(Niedenzu 1921: 14)
2.2. Ac idoca pus al e ni olius (Schumach. & Thonn.)
Nied. (Niedenzu 1915: 53)
Malpighia al e ni olia Schumach. & Thonn. (Schumache
1827: 222)
Ac idoca pus co ymbosus Hook. . (Hooke 1848: . 774)
Ac idoca pus guineensis A.Juss. (Jussieu 1840: 271)
Anomalop e ys co ymbosa (Hook. .) Kun ze (Kun ze
1891: 87)
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SUPPLEMENTARY MATERIALS
Supplemen a y ma e ial 1
GenBank accession numbe s o he species and ma ke s sampled in his s udy.
h ps://doi.o g/10.5091/plece o.158824.suppl1
Supplemen a y ma e ial 2
Mo phological ma ix wi h wo cha ac e s coded om ype and he ba ium specimens o 21 species o Ac idoca pus
and i s ou g oups.
h ps://doi.o g/10.5091/plece o.158824.suppl2
Supplemen a y ma e ial 3
In agene ic classi ica ion sys em o Ac idoca pus p oposed by Niedenzu (1928) and modi ied by A ènes (1945) and
Mo on (1968).
h ps://doi.o g/10.5091/plece o.158824.suppl3
Supplemen a y ma e ial 4
Ch onog am esul ed om he BEAST analysis, showing 95% clade c edibili y in e al o each node.
h ps://doi.o g/10.5091/plece o.158824.suppl4
