Biodi e si y Da a Jou nal 13: e161978
doi: 10.3897/BDJ.13.e161978
Taxonomy & In en o ies
Fi s eco d o Diplo axis kohlaanensis
(B assicaceae) o he eas e n A ican lo a wi h
axonomical and phy ogeog aphical ema ks
Sal a o e Camb ia , Manuela Po o ecchio , Ad iana San anello , Pie o Minissale ,
Gian Pie o Giusso del Galdo
‡ Depa men o Biological, Geological and En i onmen al Science, Uni e si y o Ca ania, Via A. Longo 19, I - 95125,
Ca ania, I aly
Co esponding au ho : Sal a o e Camb ia (camb ia_sal a o [email protected] ), Pie o Minissale (p.minissale@unic .i )
Academic edi o : Giannian onio Domina
Recei ed: 13 Jun 2025 | Accep ed: 03 Aug 2025 | Published: 07 Oc 2025
Ci a ion: Camb ia S, Po o ecchio M, San anello A, Minissale P, Giusso del Galdo GP (2025) Fi s eco d o
Diplo axis kohlaanensis (B assicaceae) o he eas e n A ican lo a wi h axonomical and phy ogeog aphical
ema ks. Biodi e si y Da a Jou nal 13: e161978. h ps://doi.o g/10.3897/BDJ.13.e161978
Abs ac
Backg ound
Diplo axis (B assicaceae) is a genus widely dis ibu ed in he empe a e a eas o Eu ope,
Asia and A ica, eaching i s g ea es di e si y in NW A ica, Cape Ve de and he
Medi e anean a ea. In eas e n A ica, his genus is only ep esen ed by D. ha a
(Fo ssk.) Boiss. in Somalia and Djibou i, while no na i e species o Diplo axis ha e been
eco ded om E hiopia.
New in o ma ion
The inding o Diplo axis kohlaanensis A.G. Mill. & J.A. Nybe g, o iginally desc ibed om
no he n Yemen, is he e epo ed o he i s ime in he Simien Moun ains. Ou inding
has a pa icula phy ogeog aphical and ecological signi icance, highligh ing he ele an
lo is ic ela ionships o he A abian Peninsula moun ains wi h he E hiopian highlands. In
‡ ‡ ‡ ‡
‡
© Camb ia S e al. 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.
addi ion, ou in es iga ions con i m he ou s anding numbe o exclusi e axa sha ed by
hese wo a eas es i ying o he pas phy ogeog aphical connec ion be ween hese wo
e i o ies. D. kohlaanensis is localised on he high-al i ude cli s, which ep esen a highly
conse a i e en i onmen o elic species. Finally, he axonomical ela ionships o his
axon wi hin he Diplo axis ha a complex a e also examined.
Keywo ds
biogeog aphy, chasmophilous ege a ion, E hiopia, mo phology, axonomy
In oduc ion
Diplo axis DC., a genus o B assicaceae Bu n. ( ibe B assiceae DC.), is mainly
dis ibu ed in he empe a e a eas o Eu ope, Asia and A ica, including 30-40 species
(Ma inez-Labo de 1988, Wa wick and Hall 2009, Pignone and Ma ínez-Labo de 2010,
Gómez-Campo 2011, POWO 2025) wi h he g ea es di e si y in NW A ica, Cape Ve de
and he Medi e anean a ea (Ma ín and Sánchez-Yélamo 2000, Rus an 2008).
Acco ding o he Flo a o E hiopia and E i ea (Edwa ds e al. 2000), he mos ecen
academic lo a dealing wi h hese coun ies, no species belonging o his genus occu s in
E hiopia, while o he sou ces (Oli e 1868, Pignone and Ma ínez-Labo de 2010, POWO
2025) epo he p esence o weedy species D. e ucoides L. Fo he emaining eas e n
A ican coun ies, he only known na i e species o Diplo axis is D. ha a (Fo ssk.) Boiss.,
occu ing in no he n Somalia and Djibou i (Thulin 1993). This species, o iginally placed
by Schulz (1919) in he sec ion Ca oca pum, shows a he unce ain ela ionships and a
qui e isola ed posi ion wi hin he genus Diplo axis (P adhan e al. 1992, Wa wick e al.
1992, Ma ín and Sánchez-Yélamo (2000). Besides, his axon is mo phologically e y
a iable (Ma ínez-Labo de 1991) and widely dis ibu ed om he (sub-)dese ic
e i o ies o No h A ica eas wa ds o A ghanis an and Pakis an (Zoha y 1966, Po ie -
Alape i e 1979, Fennane 1999, POWO 2025). Two closely-allied axa, D. c assi olia (Ra .)
DC. and D. lagascana DC., some imes ea ed a subspeci ic le el, a e also known om
Sicily (I aly) and sou he n Spain, espec i ely (Cas o iejo e al. 1993, Pigna i e al. 2019,
B ullo and B ullo 2020). Besides, he pe ennial popula ions o D. ha a om No h A ica,
al e na i ely e e ed by a ious au ho s (Mai e 1965, Po ie -Alape i e 1979, Fennane
1999) o he subsp. c assi olia and lagascana, bu clea ly dis inguished by he
mo phology o he lea es and siliques, a e p o isionally a ibu ed o he a iabili y o he
species, al hough u he axonomic, ecological and cho ological in es iga ions a e
desi able. The D. ha a complex includes all he endemic axa eco ded om Cape Ve de
(Rus an 2008), whose o igin is qui e ecen (Qua e na y) likely om he wes e n Saha an
popula ions o D. ha a as p oposed by F anzke e al. (2017). Rega ding he sou he n
bounda ies o D. ha a, i eaches he cen al-sou he n pa o he A abian Peninsula and,
pa icula ly, Saudi A abia, Yemen and Oman (Mille and Cope 1996), whe e Mille and
Nybe g (1994) ound and desc ibed a new closely- ela ed species named D.
kohlaanensis om he moun ain a ea o Kuhlan (no he n Yemen). Acco ding o hese
au ho s, his axon is mo phologically and ecologically qui e dis inc om D. ha a,
2Camb ia S e al
especially o i s sh ubby habi , co olla wi h longe pe als and la ge siliques. Whe eas,
ela ing o i s ecology, i occu s on ca bona ic o calca eni ic ocks loca ed a high
al i udes (2,300-3,000 m a.s.l.), ea u ing cool and humid en i onmen al condi ions whe e
D. ha a is ypically ound in he A abian Peninsula. Recen ly, D. kohlaanensis was ound
by B inkmann e al. (2009) also in he Jabal al Akhda moun ain ange (Oman), al hough
his eco d should be conside ed a leas dubious as he ecological con ex seems
unlikely o he species. Howe e , o he epo s o Oman, such as hose o he specimens
in he GBIF da abase om he Dho a Moun ains, seem o demons a e he p esence o
he species in ha coun y.
Du ing ield su eys on he A omon ane and A oalpine ege a ion o he Simien M s.
(no h-wes e n E hiopia), a popula ion o an unknown species belonging o he genus
Diplo axis was ound on he e ical cli s jus below he peak o M . Ina ye, a an al i ude
be ween 3,700 and 3,900 m a.s.l. Fu he ield and he ba ium in es iga ions allowed us
o a ibu e his plan o he a o emen ioned D. kohlaanensis due o he peculia
mo phological ea u es o he s ems, lea es, lowe s and siliques. Ou inding is qui e
ele an since i ep esen s he i s eco d o a na i e species o he genus Diplo axis in
E hiopia, hus con ibu ing o upda e he al eady signi ican pool o species sha ed by he
E hiopian highlands and he moun ains o he sou he n A abian Peninsula. This ela i ely
su p ising disco e y in a e i o y ha is qui e well known om a lo is ic poin o iew, a
leas in he highe a eas (Pu and Nemomissa 2001, Pu and Nemomissa 2005), is
p obably ela ed o he localisa ion o he species on e ical walls ha a e di icul o
access and pe haps also o i s lowe ing pe iod in he d y season. A de ailed
mo phological desc ip ion o he plan s belonging o he su eyed popula ion, as well as
he phenology, conse a ion, dis ibu ion and ecology o he species in E hiopia is
discussed in his pape . Fu he mo e, a axonomic e iew o he cu en ly ecognised axa
wi hin he D. ha a species complex is p esen ed wi h he elabo a ion o a key. Finally, he
lo is ic ela ionships be ween he Simien Moun ains and he moun ain anges o he
sou he n A abian Peninsula a e analysed in de ail, highligh ing he ege a ion ypes ha
hos he g ea es numbe o sha ed species be ween eas e n A ica and sou h-wes e n
Asia.
Ma e ials and me hods
The s udy a ea is localised in he no h-wes e n pa o E hiopia and, in pa icula , in he
Simien M s. (13°140' N, 38°210' E) whe e he highes peak o E hiopia is ound (M . Ras
Dejen, 4,540 m a.s.l.). This massi is he emnan o a majo Oligo-Miocene shield
olcano, deeply e oded by he Tekeze Ri e and i s ibu a ies, su ounding he massi
(Kie e e al. 2004). A peculia ea u e is he p esence o a s eep slope o ca. 1,000 m
connec ing he high pla eau o he lowlands.
The a eas exceeding 3,500 m o al i ude a e cha ac e ised by a moun ain clima e wi h
equen os and occasional snow (Hu ni 1988), wi h mean daily empe a u e anging
om 1.5°C in he coldes day up o 14.6°C in he ho es day (Hu ni and S ähli 1982).
Rain alls chie ly occu du ing he we season (May-Augus ), ollowed by a d y season,
Fi s eco d o Diplo axis kohlaanensis (B assicaceae) o he eas e n ... 3
wi h mean annual alues o 1,515 mm a Gich Camp (Hu ni and S ähli 1982). Mo ing
om he no he n o he sou he n slopes o he Simien M s., ain alls a e ypically
dec easing.
Ou ield su eys we e pe o med in Decembe 2024 and Janua y 2025. Flowe ing and
ui ing specimens o Diplo axis kohlaanensis we e collec ed on he e ical cli s below
M . Ina ye (Fig. 1). As i is loca ed in a he inaccessible s ands, i was decided o collec a
limi ed quan i y o specimens by ha es ing only he elemen s ele an o he p ope
iden i ica ion o he plan , such as basal and cauline lea es, lowe s and siliques, also o
a oiding any po en ial damage o he popula ion. The collec ed specimens a e p ese ed
a he He ba ium o Ca ania (CAT, he ba ium ac onym ollows Thie s (2025)).
The mo phological in es iga ions on he E hiopian ma e ial belonging o D. kohlaanensis
we e ca ied ou on i e indi iduals. All he specimens we e compa ed wi h da a om he
p o ologue (Mille and Nybe g 1994) and o he ele an li e a u e (Zoha y 1966, Po ie -
Alape i e 1979, Thulin 1993, Mille and Cope 1996, Pigna i e al. 2019, B ullo and B ullo
2020).
Figu e 1.
O e all ange o Diplo axis kohlaanensis acco ding o li e a u e and he ba ium da a (g een
do s) and ield su eys (pu ple do ) and dis ibu ion map o D. kohlaanensis in E hiopia ( ed
do ).
4Camb ia S e al
The compa ison amongs he specimens belonging o D. ha a, D. c assi olia, D.
lagascana and D. kohlaanensis was ca ied ou using bo h li ing ma e ial and d ied
specimens p ese ed in B, BR, CAT, E, K, LI, MA, PAL, SANT, SAV, US, W and WAG. In
pa icula , he ou species ha e been di e en ia ed, based on he ollowing cha ac e s:
habi us, lea and s em indumen um, lea blade shape and size, lea ma gins, co olla
colou , pe al size and silique size and shape (including beak leng h). In addi ion, he
conse a ion s a us o D. kohlaanensis in E hiopia was assessed by using he IUCN
c i e ia (IUCN 2001) and he guidelines o egional applica ion (IUCN 2003), acco ding
o he GeoCAT (Geospa ial Conse a ion Assessmen Tool) p og amme (Bachman e al.
2011). Finally, o he phy ogeog aphic s udy, a checklis o he species occu ing in he
A omon ane and A oalpine a eas o he Simien M s. exceeding 3,000 m a.s.l. was
elabo a ed by using li e a u e (Pu and Nemomissa 2005, Melese e al. 2018) and ield
su eys. Then, wi h he suppo o he POWO da abase, i was possible o iden i y species
occu ing in he Simien M s. wi h a disjunc dis ibu ion ange in eas e n A ica and he
A abian Peninsula. Subsequen ly, o each species, he ege a ion ype was iden i ied
wi h he suppo o li e a u e (Edwa ds and Hedbe g 2009) and ield da a. Besides, he
plan li e o m o hese axa has also been in es iga ed. To compa e he plan li e o ms o
axa wi h an Eas A ica and A abian dis ibu ion, a s acked ba cha was gene a ed
using RS udio (RS udio Team 2020) and he idy and ggplo 2 lib a ies (Wickham 2016,
Wickham e al. 2017).
Da a esou ces
The ollowing he ba ium specimens ha e been examined: Diplo axis ha a (Fo ssk.)
Boiss.: YEMEN: Go . Hadh amou , Jol Pla eau, on he pipeline d. om Wadi A a o
Tawila ields, immedia ely S o he highes poin and wa e shed, 15°09'53.8"N,
49°22'37.1"E, 3 Sep embe 2001, 1620 m al ., N. Kilian, P. Hein & M. A. Hubaishan s.n.
(MA917618; WAG1970184!); OMAN: Dho a , Mughsayl, olcanic esca pmen s and
oo hills, 16.891781 N 53.823051 E, 28 Sep embe 2015, P. Escoba Ga cia s.n.
(W201602241!); Dho a , Shaa , Rass Saij Cli s, o es and clea ings, 16.763547 N
53.602058 E, 28 Sep embe 2015, P. Escoba Ga cia s.n. (W201602242!); Dho a ,
Taquah, Way o Wadi Da ba , deciduous o es , 17.070599 N 54.446298 E, 27 Sep embe
2015, P. Escoba Ga cia s.n. (W201602243!); Dho a , Al Mughsayl, Wadi in ondo alla
discesa, dopo Al Mughsayl, lungo le pendici e il le o di sinis a. Al . 50-100 m, 10
Sep embe 2002, M. Ra aelli, M. Ta deli, S. Mos i 851 (E00983905!); Dho a , Wadi
Mughsayl, al . 0-250 m, 4 Ma ch 1994, I. McLeish 3437 (E00121242); Dho a , oad om
Taqa o Medina al Haq, in ough g ass a he oad side, 500 m al ., 17,073 N 54,255 E, 7
Oc obe 1984, R.A. Ash 185 (E00449433!); JORDAN: Elgi p ope Pe am, Wadi Musa,
A abia Pe aea, 18 June 1909, F. Nàbělek 1403 (SAV0004618!); TUNISIA: Sousse, ca. 7
km on he oad Knais o Gabghoub, o - oad o a i icial lake. Open e ain wi h he bs and
sh ubs, along lake, 90 m al ., 35°40.3' N, 10°26.3' E, 2 Ap il 2003, J.J. Wie inga 4848
(BR0000025202021!); MOROCCO: G ea A las, Go ges du Ziz be ween Rich and E -
Rachidia, su oundings o Tunnel du Leginai es, 21.1 km S Rich, 1280 m al ., 31°10' N,
04°23' W, 29 Ap il 1993, R. Vog 10366 & C. Obe p iele 4814 (B100354760!); SOMALIA:
Fi s eco d o Diplo axis kohlaanensis (B assicaceae) o he eas e n ... 5
Togdhee , Bu ao Dis ic , Gaan Libaah (Gacan Libaax), ca. 140 km om Ha gyesa on
Ha gyesa-Bu an, ields, 2000 m al ., 28 Decembe 1977, K. Elmi, M. Suliman 216
(WAG0338919!); Diplo axis c assi olia (Ra in.) DC. : SICILY (ITALY): Sizilien:
Bachschluch hin e Po o Empedo le südwes l. Ag igen o, ech e Talsei e nahe dem
Ausgang, 15 Ap il 1965, 1620 m al ., H.E. Me lesics s.n. (LI253868!); Pa e nò sulle upi
calca ee ol e il Sime o, 1 May 1896, F. To nabene s.n. (CAT3970!); Pa e nò, 15 Ap ile
1894, P. Bacca ini s.n. (CAT3971!); Pie ape zia, s.d., s.c. (PAL5624!); To e di Ga e,
Lica a (AG), 25 June 1991, G. Ce a, F. Gendusa, E. Pi a s.n. (PAL71359!); Diplo axis
lagascana DC.: SPAIN: Alme ía: Tabe nas, Llanos del Duque, sob e subs a o seco,
ped egoso y salino, 20 Ma ch 1998, C.Mo ales, C.Quesada, L.Baena, J.E.Lina es s.n.
(SANT 40196!); Velez de Benaudalla (Espana, p o . G anada, Andalucia), al . 50 m,
campos de cul i o al lado de la ca e e a, comunidade de e o i os, 17 Ma ch 1982, A.M.
Neg illo, P. A oza, J. Hu ado 12186 (BR000027543092V!); Velez de Benaudalla
(Espagne, p o . G anada, Andalucia), El Azud de Vèlez, 40 m al ., bo ds des chemins,
su des sols emuès iches en pe i es pie es, 28 Janua y 1980, M. Lade o, O. Soco o, J.
Hu ado 10240 (BR000026282237!); Base de Sie a G ossa, Alican e, 11 No embe
1949, S. Ri as s.n. (US03584685!); Diplo axis kohlaanensis A.G. Mill. & J.A. Nybe g:
YEMEN: Am an o Kuhlan Road, limes one cli s, 15 km E o Kuhlan, 2800 m al ., 26
Ma ch 1981, A.G. Mille & D.G. Long 3213 (E0038048! holo ype; K001291634! iso ype);
on cli s abo e he own o Kohlaan, 2700 m al ., 31 Janua y 1979, J.R.I. Wood 2691,
iden i ied by A. G. Mille (E00449402!); on a cli s abo e Kohlaan, 2300 m al ., 17
Feb ua y 1973, J.R.I. Wood 2691, iden i ied by A. G. Mille (E00449405!); on cli s on he
S Wes edge o Jebel Mahdad, 3000 m al ., 26 Sep embe 1978, J.R.I. Wood 2518,
iden i ied by A. G. Mille (E00449403!); Bai Al Alama, J. Al Mahdad, 20 km W o Am an,
SW acing sands one cli s, 2900 m al ., 26 Sep embe 1978, A.G. Mille 236
(E00449404!); OMAN: Wes e n Haja Moun ains, s eep N-Exposed cli in Teuc io-
Junipe e um, 16 Ap il 2011, A. Pa zel s.n. (E00702050!); ETHIOPIA: Simien M ., ocky
cli s nea he pa h be ween Ina ye Moun ain and Chennek, 3900 m al ., 13°15'20.17"N,
38°10'9.31"E, 23 Decembe 2024, S. Camb ia s.n. (CAT!).
Taxon ea men
Diplo axis kohlaanensis A.G. Mille & J.A. Nybe g 1994
Ma e ial
a. scien i icNameID: Diplo axis kohlaeensis; kingdom: Plan ae; o de : B assicales; amily:
B assicaceae; genus: Diplo axis; con inen : A ica; coun y: E hiopia; locali y: Simien
Moun ains; e ba imEle a ion: 3900 m; e ba imLa i ude: 13°15'20.17"N;
e ba imLongi ude: 2 38°10'9.31"E; indi idualCoun : 100; iden i iedBy: S. Camb ia;
da eIden i ied: 23-12-2024; occu enceID: 0DD55C97-CA41-5778-B3F7-
F8CECAA94124
6Camb ia S e al
Desc ip ion
Sh ubby and compac chamaephy e, gene ally glab ous, wi h many s ems a ising
om a woody oo s ock. Flowe ing b anches e ec o ascending, 30–60 cm long.
Lea es sligh ly leshy, glaucous, all pe iola e wi h a well-ma ked whi ish mid- ib. The
lowe ones wi h o a e o oblong-o a e blade, 15–80 × 10–20 cm, ob use apex and
se a e ma gin wi h 4–6 pai s o ee h o sinua e o en i e, a ely wi h ew sca e ed
hai s. Uppe lea es smalle , 5–40 × 5–10 mm, p o ided wi h a sho e pe iole.
Racemose in lo escence, e ec o ascending, wi h he lowe s o e opping he buds.
Flowe pedicels 10–14 mm long, equal o sho e han o he pe als. Calyx wi h
ou
sepals, 5–6 mm long, gene ally 2.5–3 imes sho e han he pe als, ex e nally hai y
pa icula ly along he mid- ib and nea he base. Inne sepals 1.8-2.5 mm b oad and
sacca e a base, he ou e 1.5–2 mm b oad, wi h non-sacca e base and clea ly
hooded ip. Co olla wi h ou pe als, pale yellow, b oadly obo a e and na owing
below in o a linea claw, 10–14 × 5-6 mm, wi h ounded ip. And oecium consis ing o
6 s amens; he median 8–11 mm long, wi h p ominen nec a ial glands a he base;
he la e al 6–7 mm; an he s 2–3 mm long. Gynoecium wi h cylind ical o a y, glab ous,
sho ly s ipi a e; s igma bilobed. Siliques e ec o pa en , linea o linea -oblong,
la ened, 12–40 × 2.5–3.2 mm, wi h he seeds in wo ows; beak seedless, 1.5–2 mm
long; Seeds pale eddish o b own, la ened, 1–1.5 × 0.5–l mm (Fig. 2, Fig. 3).
Figu e 2.
Diplo axis kohlaanensis in he E hiopian s ands: A Na u al habi a whe e an indi idual is
highligh ed by a ed ci cle; B Chasmophilous ege a ion wi h Aeonium leucoblepha um and D.
kohlaanensis; C Habi us o D. kohlaanensis; D De ail o he lea es and lowe s.
Fi s eco d o Diplo axis kohlaanensis (B assicaceae) o he eas e n ... 7
Dis ibu ion
Based on he cu en knowledge, he species is localised in E hiopia on he e ical,
basal ic, no h- acing cli s o he Simien M s. a an al i ude anging om 3,700 o
3,900 m a.s.l.
Ecology
The species was ound on he esca pmen below M . Ina ye, which is cha ac e ised
by
chasmophy ic ege a ion ich in a e and phy ogeog aphically ele an species, such
as Aeonium leucoblepha um Webb ex A.Rich., A abis alpina L. subsp. alpina,
Campanula edulis Fo ssk , Poa simensis Hochs . ex A.Rich., Helich ysum ci ispinum
Delile a . ci ispinum, Dian hus longiglumis Delile, Rosula ia semiensis (J.Gay ex
A.Rich.) H.Ohba, Asplenium ae hiopicum (Bu m. .) Beche e and ma ginally also
Figu e 3.
Iconog aphy o Diplo axis kohlaanensis, based on ma e ial om E hiopia. D awing by Ad iana
San anello.
8Camb ia S e al
P imula e icilla a Hochs . subsp. simensis (Hochs .) W.W.Sm. & Fo es [=E o ochis
simensis (Hochs .) Fı a & Lidén].
Conse a ion
The popula ion o D. kohlaanensis en i ely alls wi hin he Simien Na ional Pa k, a
p o ec ed a ea c ea ed in 1969, a pe iod when 80% o he Pa k was subjec ed o
human exploi a ion h ough li es ock g azing, cul i a ion and se lemen s (Debonne
e al. 2006). Ac ually, his is s ill h ea ened by human ac i i ies, pa icula ly g azing
and logging (Jacob e al. 2016), u he agg a a ed by he ecen wa e en s which
led o a se e e educ ion o he wooded a eas due o coppicing and i es, as
highligh ed by Meaza e al. (2025) o o he moun ains o no he n E hiopia. Fo his
eason, he Simien Moun ain Na ional Pa k has been lis ed as a Wo ld He i age Si e
in Dange since 1997. Despi e his, he e ical cli s a e poo ly a ec ed by human
dis u bance due o hei inaccessibili y and, he e o e, hey ep esen he bes -
p ese ed en i onmen s o his a ea. The popula ion o D. kohlaanensis con ains a
e y low numbe o indi iduals ( oughly es ima ed in ca. 100 ma u e plan s) ha , e en
i no immedia ely h ea ened, a his s age, we do no know how he species would
espond o global wa ning. By using he Geospa ial Conse a ion Assessmen Tool
(Bachman e al. 2011), he ex en o occu ence (EOO) is calcula ed o be 0.028 km ,
while he a ea o occupancy (AOO) is 4 km . Following he IUCN C i e ia a global
and egional le el (IUCN 2001, IUCN 2003), wi h he es ima ed EOO less han 100
km , an AOO less han 10 km , he conse a ion s a us o D. kohlaanensis o E hiopia
has been classi ied as Endange ed a egional le el (EN eg: D) acco ding o c i e ion
D.
Biology
Flowe ing and ui ing du ing he d y season om No embe o Ma ch.
Iden i ica ion keys
Key o he axa belonging o Diplo axis ha a species complex
1Annual o a ely pe ennial he b, loosely b anched, wi h densely hai y lea es
and s ems, lea es no leshy wi h 5–15 pai s o ee h Diplo axis ha a
–
Pe ennial su u icose plan , wi h a basal woody oo s ock, densely b anched,
wi h glab ous o glab escen lea es and s ems, lea es leshy wi h 1–6 pai s
o ee h o pinna ipa i e
2
2
Flowe ing b anches lexuosus, lea es wi h den a e o lobed ma gins, pe als
7–9 mm long, sho e han lowe pedicel, da k yellow, siliques hanging a
ui ing, lowe s no o e opping he buds
3
2
2
2 2
Fi s eco d o Diplo axis kohlaanensis (B assicaceae) o he eas e n ... 9
D. ha a D. lagascana D. c assi olia D. kohlaanensis
Flowe ing b anches
leng h (cm)
60 80 100 300
In lo escence e ec e ec lexuous and hanging e ec
Pe al colou da k yellow da k yellow da k yellow pale yellow
Pe al leng h (mm) 7–9 7–9 6–10 10–14
Siliques size (mm) 10–50 × 2–3 25-40 × 1.5–3.5 30–60 × 2–3 12–40 × 2.5–3.2
Beak leng h (mm) 1.5–2 1–2 1–2 1.5–2
The wo Eu opean species o he D. ha a complex, namely D. c assi olia and D.
lagascana, show a g ea e mo phological a ini y wi h D. kohlaanensis, especially o he
subsh ubby habi and he glab ous (o glab escen ) and leshy lea es. D. lagascana, a
Spanish endemism o en ea ed a subspeci ic ank (sub D. ha a (Fo ssk.) Boiss. subsp.
lagascana (DC.) O.Bolòs & Vigo) is easily dis inguishable o ha ing pinna ipa i e o
pinna i idous lea es (Cas o iejo e al. 1993). As conce ns D. c assi olia [=D. ha a
(Fo ssk.) Boiss. subsp. c assi olia (Ra .) Mai e], a Sicilian endemism (Pigna i e al. 2019,
B ullo and B ullo 2020), w ongly eco ded also o No h A ica (Mai e 1965, Po ie -
Figu e 5.
Main mo phological ea u es o he axa belonging o he Diplo axis ha a g oup - lowe ing
b anches (1); Lea (2); Silique (3): A D. kohlaanensis; B D. c assi olia; C D. lagascana; D D.
ha a. D awings by Rosa ia Di Cicca.
16 Camb ia S e al
Alape i e 1979, Fennane 1999), shows a simila lea blade, o en den a e wi h 1–6 ee h
o side. Howe e , i is easily dis inguished om D. kohlaanensis by i s long, lexible and
pendulous lowe ing b anches and by i s lowe s wi h da k yellow and much sho e
pe als (6-10 mm), always sho e han he peduncles. In conclusion, he species a issue
is e y well dis inguishable om he o he axa o he D. ha a g oup, no only o i s
mo phology, bu also o i s ecological beha iou and geog aphical dis ibu ion.
The e o e, he disco e y o his specie in E hiopia has a ema kable phy ogeog aphic
alue, since i is he i s eco d o a Diplo axis species in E hiopia and, likely mos
in e es ing, con i ms he phy ogeog aphic ela ionships be ween he E hiopian highlands
and he A abian Peninsula moun ains.
Ou in es iga ions on he A omon ane and A oalpine species wi h an Eas A ican-
A abian dis ibu ion (i.e. amphi-Red Sea axa) occu ing in he Simien M s. highligh he
clea p e alence o pe ennial, he baceous o woody species, mainly linked o he
A omon ane g asslands and woodlands. This is due o he lack o moun ains in he
A abian Peninsula highe han 3,600 m a.s.l., no allowing he p esence o a signi ican
A oalpine lo a ypical o he highes peaks o he Simien M s.. The e o e, he moun ain
lo a in common is mos ly ep esen ed in he al i udinal bel co e ed by E ica woods,
sh ubs and seconda y g asslands. Con e sely, he signi ican occu ence o
chamaephy es mus be highligh ed showing his dis ibu ion in e ical cli s e en a
al i udes ypical o he A oalpine bel . In ac , his en i onmen ep esen s a well-known
conse a i e and s able habi a (La son e al. 2000, Ga cía e al. 2020, Sangüesa‐
Ba eda e al. 2022, Múgica Ca nice o e al. 2024), whe e, due o he mino human
dis u bance and he peculia mic oclima e, ancien species, o en endemic, ha e been
p ese ed, p o iding p ecious e idence o a lo a which ha e now la gely disappea ed in
he nea by a eas. In his pe spec i e, Diplo axis kohlaanensis can be in e p e ed as a
elic species, linked o pa icula ly cool and humid mic oclima e ypical o high-al i ude
no h-exposed cli s. Fu he mo e, i shows some a chaic ea u es wi hin he Diplo axis
genus, such as he woody habi (Lems 1960, B amwell 1972, C onk 2006, C onk 2008,
Lens e al. 2013), sugges ing a po en ial ances y compa ed o some allied
Medi e anean he baceous species, as al eady highligh ed by Pigna i (1979) and by
B amwell (1976) o some Maca onesian-A ican axa B amwell (1976). Ac ually, he
phy ogeog aphical a ini ies be ween he Medi e anean a ea and he Ho n o A ica ha e
al eady been highligh ed by se e al au ho s (Bal ou 1888, Gille 1941, La anos 1975,
Fici 1991), pa icula ly o he no he n Somali esca pmen , an a ea wi h se e al axa
ha ing close Medi e anean allies. Acco ding o Wickens (1975), he sou he n A abian
Peninsula and eas e n A ica ep esen ed an impo an cen e o e uge, specia ion and
adia ion o Hola c ic axa du ing he glacial phases o Qua e na y. E en oday, hese
species s ongly cha ac e ise he lo a o he E hiopian pla eau, which, compa ed o o he
moun ain anges o eas e n A ica (Hedbe g 1957, Hedbe g 1964, Hedbe g 1969,
Bussmann 2006, Camb ia e al. 2024) shows a lowe incidence o he ypical A oalpine
lo a and a mo e signi ican p esence o he Hola c ic elemen . In his ame, he disco e y
o D. kohlaanensis sugges s he ole o he high al i ude cli s o he Siemen M s. as a
e uge habi a o se e al species wi h no he n a ini ies, whose o igin p obably da e
back o he glacial phases o Pleis ocene when, as a esul o a ela i ely cool and humid
Fi s eco d o Diplo axis kohlaanensis (B assicaceae) o he eas e n ... 17
clima e, nume ous species mig a ed om he no h h ough he Red Sea co ido
eaching he A abian Peninsula and he Ho n o A ica. This o igin can be pos ula ed also
o o he species g owing in he same upicolous habi a o D. kohlaanensis, such as
Dian hus longiglumis Delile, P imula e icilla e Fo ssk., S achys icha diana R.K . Singh,
Sonchus melanolepis F esen, A abis alpina L. e c. As conce ns Aeonium
leucoblepha um Webb ex A.Rich., equen in he same en i onmen s, i s ma ked
geog aphical isola ion om all o he species o i s genus should be conside ed as he
e ec o a long-dis ance dispe sal du ing he Pleis ocene, a he han as he esul o
olde ancien phy ogeog aphic connec ions be ween he Maca onesian Islands and Eas
A ica (Mo e al. 2002). Fu he esea ch in he Siemen M s. esca pmen may lead o
o he impo an lo is ic indings, also complemen ed by he mos ecen echnological
ools such as d ones, which ha e p o en pa icula ly use ul in he in es iga ion o simila
inaccessible e ical cli s (Po o ecchio e al. 2024, Ta illa e al. 2024, Wagne e al.
2024). Likewise, phylogene ic s udies aiming a cla i ying he ela ionships be ween D.
kohlaanensis and he o he axa o D. ha a complex a e highly desi able.
Acknowledgemen s
This in es iga ion was inancially suppo ed by he esea ch p og amme (PIA.CE.RI.
2024-2026 cod. 22722132189) unded by he Uni e si y o Ca ania. We hank Rosa ia Di
Cicca o he ealisa ion o he iconog aphies o Fig. 5.
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