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Hitherto overlooked or a recent introduction? (Re)discovery of Ornithopus perpusillus in Hungary

Author: Mesterházy, Attila; Bauer, Norbert; Király, Gergely
Publisher: Zenodo
DOI: 10.33542/TJB2025-2-02
Source: https://zenodo.org/records/17734068/files/02_Mesterhazy-et-al._final_48-56.pdf
48
Thaiszia - J. Bo ., Košice, 35: 048–056, 2025
h ps://doi.o g/10.33542/TJB2025-2-02
THAISZIA
JOURNAL OF BOTANY
Hi he o o e looked o a ecen in oduc ion? (Re)disco e y o
O ni hopus pe pusillus in Hunga y
A ila Mes e házy1, No be Baue 2 & Ge gely Ki ály3
1 H-9500, Hunyadi u ca 55., Celldömölk, Hunga y; [email p o ec ed]
2 Depa men o Bo any, HNM Public Collec ion Cen e, Hunga ian Na u al His o y Museum, H-1087
Budapes , Köny es Kálmán k . 40, Hunga y; [email p o ec ed]
3 Uni e si y o Sop on, Facul y o Fo es y, H-9400, Bajcsy-Zs. u. 4., Sop on, Hunga y;
kbge [email protected]
Mes e házy A., Baue N. & Ki ály G. (2025): Hi he o o e looked o a
ecen in oduc ion? (Re)disco e y o O ni hopus pe pusillus in
Hunga y. – Thaiszia – J. Bo . 35: 048–056.
Abs ac : This a icle p esen s he hi he o known eco ds o O ni hopus
pe pusillus in Hunga y. The species was i s ound in cen al Hunga y as
ea ly as 1952, bu was iden i ied as O. sa i us. I was ( e-)disco e ed in
sou hwes e n Hunga y in 2010 a se e al locali ies, in dis u bed, open
sandy g asslands, wi h co-occu ence o o he A lan ic-Medi e anean
species. O. pe pusillus was likely in oduced in Hunga y a he beginning
o 2000s, and now i is conside ed o be na u alized he e.
Keywo ds: Cen al-Eu ope, sandy ege a ion, Leguminosae, in oduced
species.
In oduc ion
The genus O ni hopus L. (Leguminosae), comp ising i e species (O ni hopus
sa i us B o ., O. comp essus L., O. mic an hus (Ben h.) A ., O. pinna us (Mille )
D uce., and O. pe pusillus) occu mainly in he wes e n Medi e anean and in No h
A ica. Only one species (O. mic an hus) is na i e in Sou h Ame ica (Polhill 1981). The
species o he genus a e annuals wi h small pink o yellow pe als, he ui b eaks
along ans e se su u es be ween seeds du ing ipening. The Eu opean species o
he genus can be ound mainly in dis u bed acidophilous sandy g asslands o in
ude al ege a ion (Glads ones & Ba e -Lenna d 1964). Th ee species (O.
49
comp essus, O. pinna us, O. sa i us) ha e been cul i a ed p e iously as odde plan s
ou side hei na u al ange (No h Ame ica, Eas e n Eu ope, Aus alia, New Zealand).
Hunga ian lo as o he las hal cen u y (Soó 1964; Simon 2000; Ki ály 2009)
epo ed only O. sa i us in Hunga y; his species was o me ly cul i a ed on nu ien -
poo soils, mainly wi h ce eals (Csapody & P isz e 1961).
Ma e ial and Me hods
Geog aphical coo dina es we e eco ded using handhold GPS de ice. Mapping
uni s we e de ined acco ding o he Cen al Eu opean Flo a Mapping P og amme
(Nikl eld 1971). Nomencla u e o plan names ollows he Eu oMed (2006-), while
names o plan associa ions ollow Bo hidi e al. (2012). Fo he small geog aphical
egions o Hunga y, we elied on he wo k o Dö ényi (2010). Du ing ou esea ch,
we e iewed all o BP's O ni hopus ma e ial. Specimens collec ed du ing he s udy
a e deposi ed in BP. The ege a ion o O. pe pusillus si e a Bolhás was sampled in a
six 0.25 m2 mic o quad a s. The abundance o plan species was es ima ed isually
as a pe cen age o g ound co e in each plo . He ba ia abb e ia ions a e acco ding
o Thie s (2025).
Resul s and Discussion
Taxonomy and iden i ica ion
O ni hopus pe pusillus is o en con used wi h he e y simila O. sa i us. The i s
Hunga ian specimen was co ec ly de e mined by he o iginal collec o bu was la e
e oneously e ised o O. sa i us. The main eason o misiden i ica ion is he ole o
quan i a i e cha ac e s ( he size o he co olla and he leng h o he lowe calyx-
ee h), hese wo species canno be dis inguished well on he ba ium specimens. The
lowe s o O. sa i us a e la ge , co olla is 5–8 mm long, pink, c eam-colo ed, less
o en whi ish, i s eins a e no conspicuous. The co olla o O. pe pusillus is smalle
(3–4 mm long), whi ish o pale pink, wi h a conspicuous ein. Howe e , hese
quan i a i e cha ac e s a e eliably isible only on esh ma e ial. Some sou ces (i.e.
Ball 1968; Jäge & We ne 2005) also men ion he a io o leng h o he b ac and
in lo escence as a dis inguishing cha ac e , bu Wallnö e e al. (2015) sugges ha
his is un eliable because he leng h o b ac s can a y widely e en on he same
indi idual. Acco ding o ou obse a ions, he e a e also di e ences in legumes
be ween he wo species. F ui s o O. pe pusillus a e always g adually cu ed, while
he legumes o O. sa i us a e almos s aigh . The hai s on ui s o he o me species
a e one and a hal o wo imes longe and dense han hose o he la e , while ui
o O. sa i us seems o be b igh e and appea ing almos smoo h. As he i s species
is ce ainly p esen he Hunga ian lo a, we included i in he o me Hunga ian
O ni hopus key (Ki ály 2009).
50
O ni hopus L.
Co olla 3–4 mm long, whi e o pinkish wi h conspicuous pu ple eins, calyx ube 2–3 × longe
han calyx- ee h, lowe calyx- ee h iangula , < 1mm long. Legume g adually cu ed, wi h
dense hai s. Plan 10–30 cm high. V-VIII. Ad . Open, dis u bed sand g asslands. (Belső-
Somogy, Bakonyalja†) ………………………………………………………………………………….. O. pe pusillus L.
Co olla 5–8 mm long, pinkish o c eam-colo ed wi h inconspicuous eins, calyx ube and calyx-
ee h a e equal size, lowe calyx- ee hs elonga ed, > 1mm long. Legume ± s aigh , wi h
sca e ed hai s. Plan 30–60 cm high. VI-VIII. Ad . (DNy-Eu.) Fodde plan , o me ly cul i a ed
……………………………………………………………………………………………………………………… O. sa i us B o .
Dis ibu ion
O. pe pusillus is an A lan ic-Medi e anean species, occu ing in wes e n and
sou h-wes e n pa s o Eu ope, om NW Russia (wi h he eas e nmos occu ence
nea S . Pe e sbu g, Vasilje a 1984) o he B i ish Isles and sou he n Scandina ia
(Ingelög e al. 1993). The sou heas e n ma gin o he dis ibu ion is nea Dnip o and
Khe son in Uk aine (P okudin 1987). I is no a e in I aly and he Ibe ian Peninsula
bu sca e ed in no he n Alge ia (Wie sema e al. 1990). The species is in oduced
in Es onia (island o Hiummaa, Tabaka e al. 1996). I was ound on he Cu onian Spi ,
and in wo mo e si es in cen al Li huania (Vilkonis & S i skis 2006). Ou side Eu ope
i has also been in oduced o India, Mongolia, he Caucasus, sou h Aus alia, New
Zealand, Cos a Rica, and eas o Uni ed S a es (POWO 2025).
O. pe pusillus is a e in Cen al Eu ope. In Ge many, i is mos equen on he
sandy banks o la ge i e s (Ode , Elbe and Rhine) and qui e sca e ed in No he n
Ba a ia and Thu ingia (Ma kg a 1976). In Poland, i g ows on he sandy banks o he
la ge i e s Ode and he Vis ula, and in sandy ields nea he sea (Zając & Zając
2001). I was empo a ily in oduced on sandy pas u es and in pine o es s in Czech
Republic. I is epo ed om P ague, Vel us, Li oměřice and No á Bys řice. In 1992,
one s and was ound in a sand pi nea Rumbu k (NW Czechia) (Ch ko á 1995).
Fu he mo e, wo his o ical si es ha e ecen ly been con i med in eas e n Bohemia
nea he illages Š ěpáno sko and Zdelo (Doležal 2014). I s s a us in Czech Republic
is unce ain; i occu s in seconda y habi a s, on he o he hand, O. pe pusillus has
simila dis ibu ion a ea wi h o he sub-A lan ic species (e.g. Ca ex pseudob izoides
Cla aud, Rubus gemina us H.E. Webe , Lu onium na ans (L.) Ra .) conside ed na i e.
Acco ding o Ch ko á (1995) i is non-na i e, bu i is no lis ed in he Ca alogue o
alien plan s o he Czech Republic (Pyšek e al. 2012). G ulich (2012) classi ied i as
c i ically endange ed. I seems ha he species has ecen ly been sp eading in he
Czech Republic, wi h 16 da a poin s al eady epo ed in he la es syn hesis (Chy ý
e al. 2021)
The species is conside ed a casual neophy e in Slo akia, which appea ed in 1989
(Med ecká e al. 2012). The e is a single eco d om Malé Ka pa y M s, Zá uby
Na u e P ese e, which is, howe e , axonomically unce ain and p obably e e s o
s e ile Hippoc epis comosa (Hollá 1989).
51
In Aus ia, i was i s men ioned by Neil eich (1859) om Nußdo ( oday pa o
Vienna). Beck on Mannage a (1890) accep ed his occu ence and added wo mo e
eco ds om he a ea o Vienna. Due o he lack o he ba ium ouche s hese
eco ds a e uncon i med. The e is one collec ing labelled as „Salzbu g, on sandy soil”
wi hou da e (LI). Gi en ha he plan occu s in se e al places in nea by Ba a ia,
hese obse a ions om Salzbu g a e likely o be co ec (Wallnö e e al. 2015).
Occu ences and habi a s in Hunga y
Fi s men ion o O. pe pusillus om Hunga y appea ed in Já o ka (1925) as an
occasional alien wi hou a p ecise loca ion. The sou ce o his ecod was p obably
he specimen collec ed by Józse Má on in So okúj alu ( oday: So okpolány, W
Hunga y) “in a enosis” a he beginning o he 19 h cen u y (BP101864). On his
shee , Já o ka’s handw i ing can be ound as ollows: “ex Hunga ie? subcul u es?”.
Thus, Já o ka had doub s abou he o igin o his collec ion, as J. Má on o en
labelled specimens as So okúj alu o So ki ó alu ecei ed om o he pa s o
Hunga y o e en om ab oad. In addi ion, sandy soils a e no p esen in his egion.
Ano he specimen (BP197678) om Hunga y has been ecen ly ound a BP in he
collec ion o Miklós Szalay, a pa ish p ies om Halimba (cen al Hunga y). The exac
loca ion o collec ion is no p esen ed, only one sen ence is w i en on he label:
“escaping om a o me cul u e”. As double s o Szalay’s he ba ium a e deposi ed
in he HNHM Bakony Museum (Zi c), du ing he e iew o he collec ion we ound
ano he shee (Zi c, specimen ID: 002269), on which he de ails o he collec ion had
been cla i ied (“in p a o „Csá da é ” p . pag Halimba, Hunga ia”). The specimen had
o iginally iden i ied co ec ly as O. pe pusillus, bu in 2001, i had been e oneously
e ised and published as O. sa i us by I. Galambos (Galambos 2005). The BP
he ba ium ecei ed only a ew specimens o Szalay's collec ion; hese we e
p esumably sen o S. Já o ka o iden i ica ion. P obably, he specimen o O.
pe pusillus could ha e a i ed in Budapes in his way. “Csá da- é ” is loca ed in he
wes o Halimba, a illage a he edge o Bakony M s. La ges pa o he si e ha e
been des oyed by bauxi e mining, bu some nu ien poo meadows o med on
limes one and g a el, and small acidophilous sandy g asslands s ill exis he e.
Acco ding o ou obse a ion se e al species (Ai a ca yophyllea L., An hoxan hum
odo a um L., Hype icum humi usum L., Moenchia man ica (L.) Ba l.) wi h simila
habi a p e e ence o O. pe pusillus occu in he a ea ecen ly, he e o e Szalay’s
collec ion can be accep ed as he i s occu ence o his species in Hunga y.
The species was edisco e ed in he ea ly 2000’s in he sou he n pa o Somogy
coun y (Belső-Somogy, SW Hunga y). I has since been ound in se e al places in his
egion and is cu en ly known om he ollowing loca ions:
o Rinyabesenyő (as O. sa i us, wi hou ouche , bu mos p obably O.
pe pusillus), 2004; N. P ei e (in Ba ha e al. 2015); common in pionee
ege a ion a he edge o dus oad S o he illage (46°9'12.54"N,
17°31'10.67"E), (9871.1), 22.05.2023; M. Wol , Fig. 1.
52
o Be ween Szulok and Ba cs, allow land, common (45°59'49"N, 17°31'05"E),
(0071.1), 08.06.2020, A. Mes e házy (BP).
o Bolhás, allow ield, sca e ed (46°13'57.81"N, 17°15'52.18"E), (9769.4),
26.07.2010, A. Mes e házy.
o Bolhás, Co ynepho e um, sca e ed (46°16'40"N, 17°14'40"E), (9769.1),
30.05.2017, A. Mes e házy.
o Kaszó, dis u bed g assland 0,5 km SE o Kanizsabe ek (46°19'40"N,
17°11'59"E), (9669.3), 11.05.2011, G. Ki ály.
o Kaszó, g assland wi h pionee ege a ion unde powe line 0.7 km SE o
he se lemen (46°18'48"N, 17°13'44"E), (9669.3), 18.05.2018, G. Ki ály.
Based on ecen eco ds, he species is p esen in a ela i ely la ge a ea on open
sandy g asslands. I s occu ence is mainly connec ed o ypical Filagini-Vulpine um
habi a s o med on young allows, especially in dis u bed sandy g asslands, and
Co ynepho us g asslands wi h pionee and weed species (see Table 1).
Conside a ions on he s a us in Hunga y
Vege a ion o sandy g asslands o Belső-Somogy is well s udied (Bo os 1925; Láje
1998), and he Co ynepho us g asslands ha e ecen ly been mapped (Láje 2005).
The species composi ion o he pionee g asslands o Belső-Somogy egion is ela ed
o he sub-A lan ic acidophilous sandy g asslands (Láje 2005). Many o he species
ound he e (An hoxan um puelii Lecoq & Lamo e, Aphanes aus alis Rydb.,
Teesdalia nudicaulis (L.) W.T. Ai on, Ve onica dillenii C an z.) ha e a simila
dis ibu ion in Sou hwes e n Eu ope. As he o iginal habi a s o O. pe pusillus a e
simila o hese species (Rodwell 2000; Vilkonis & S i skis 2006), he na i eness o
his species may a ise in he sandy egion o Belső- Somogy. This egion is a ec ed
by a s ong sub-A lan ic and sub-Medi e anean clima e, wi h annual ain all o
abou 780–830 mm (Bo hidi 1958). The numbe o os y days he e is one o he
lowes in Hunga y (Va ga 2003). Thus, O. pe pusillus appea ed in his a ea as in i s
Fig. 1 Flowe and legume o O ni hopus pe pusillus (le ) and pionee ege a ion a Rinyabesenyő
( igh , all pho og aphed by M. Wol ).

53
Tab. 1 Vege a ion o O ni hopus pe pusillus s ands eco ded in % a six 0.25 m2 mic o quad a s a
Bolhás Co ynepho e um (9769.1, 30.05. 2017, A. Mes e házy, 46°16'40.36"N, 17°14'39.61"E).
1.
2.
3.
4.
5.
6.
E0 moss laye
22
16
10
35
E1 he b laye
45
40
30
25
22
52
Co ynepho us canescens
45
30
25
20
15
Teesdalia nudicaulis
1
2
Amb osia a emisii olia
4
Cynodon dac ylon
6
Plan ago lanceola a
4
Hype icum pe o a um
2
4
Jasione mon ana
8
An hoxan hum puelii
2
35
O ni hopus pe pusillus
1
2
5
Conyza canadensis
2
Po en illa a gen ea
4
Eupho bia cypa issias
1
6
An hemis a ensis
0,2
Hypochoe is adica a
2
Rumex ace osella
0,5
Filago minima
20
Scle an hus annuus
1
2
na u al ange. Seeds o Leguminosae species a e iable o a long ime (Rols on
1978), he e o e O. pe pusillus may ha e been la en in his a ea o decades. In he
pas , he dis ibu ion o open g asslands in Belső-Somogy was signi ican ly highe
han oday, his habi a ype was hea ily g azed, so he sui able mic ohabi a s o his
species we e widesp ead. Howe e , he ecen co e age o g asslands in Belső-
Somogy has declined, many open habi a s ha e been a o es ed. I is likely ha – as
An hoxan hum puelii in simila habi a s – i is in a ecen in oduc ion. A. puelii was
disco e ed in 2004 in Belső-Somogy and sp ead quickly in ex ensi e a able lands and
allows. The sou ce o i s in oduc ion has been suspec ed in he seed anks o
combine ha es e s pu chased by smallholde s om no he n Ge many in he las
wo decades (Pinke e al. 2006). In he case o O. pe pusillus, he in oduc ion by
wes e n Eu opean gues hun e s is conside ed he mos likely pa hway, as he high-
quali y popula ions o dee s in he egion has always a ac ed o eign hun e s, so
he seed o he species may ha e a i ed in he a ea wi h hei ca wheels o
oo wea . As he plan is an impo an sou ce o p o ein, we ha e obse ed ha
games ( ed dee , allow dee , wild boa ) consume i s shoo s, so his may play a ole
in i s egional dis ibu ion (endozoocho y). The No h Ame ican Hype icum mu ilum
L. was disco e ed in Lake Balá a in he egion in 2001, and he au ho s a ibu ed he
occu ence o his species o hun e s oo (P ei e & Molná 2002).
54
Acknowledgemen
Má yás Wol (Kapos á ) is hanked o p o iding his da a and he pho os o he species.
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