NAT. CROAT.
VOL. 31
No 1 133-206 ZAGREB
July 31, 2022
professional paper/stručni članak – museum collections/muzejske zbirke
DOI 10.20302/NC.2022.31.13
PLETHORA OF PLANTS – COLLECTIONS OF
THE BOTANICAL GARDEN, FACULTY
OF SCIENCE, UNIVERSITY OF ZAGREB
(7): HISTORICAL OVERVIEW OF FERN
(MONILOPHYTA; POLYPODIOPSIDA;
POLYPODIOPHYTA) COLLECTIONS
Sanja Kovačić
Botanical Garden, Department of Biology, Faculty of Science, University of Zagreb,
Marulićev trg 9a, HR-10000 Zagreb, Croatia (e-mail: sanja.kovacic@biol.pmf.hr )
Kovačić, S.: Plethora of plants – Collections of the Botanical Garden, Faculty of Science, University of Zagreb (7): Historical overview of Fern (Monilophyta; Polypodiopsida; Polypodiophyta)
Collections. Nat. Croat., Vol. 31, No. 1, 133-206, 2022, Zagreb.
This sequel provides a historical overview of collections of ferns (Monilophyta; Polypodiopsida or
Polypodiophyta) grown in the Zagreb Faculty of Science Botanical Garden between 1895 and 2021.
Although opinions on the systematics, classification, nomenclature and taxonomy of “true” ferns still
diverge greatly, Equisetaceae and Psilotaceae are commonly included. At least 376 taxa of native and
cultivated ferns from 29 families have been grown in the Botanical Garden during the last 126 years.
Unfortunately, the complete COVID-19-lockdown, devastating earthquakes and weather extremes of
2020, extending to 2021, diminished our collections. Today we are growing 86 taxa within 20 families
of ferns and their closest relatives.
Key words: Zagreb Botanical Garden, Faculty of Science, historic plant collections, ferns, horsetails,
psilotum
Kovačić, S.: Obilje bilja – zbirke Botaničkoga vrta Prirodoslovno-matematičkog fakulteta
Sveučilišta u Zagrebu (7): Povijesni pregled zbirki papratnjača (Monilophyta; Polypodiopsida; Polypodiophyta). Nat. Croat., Vol. 31, No. 1, 133-206, 2022, Zagreb.
U ovom nastavku serije o zbirkama Botaničkog vrta PMF-a u Zagrebu donosim povijesni prikaz
zbirki papratnjača (Monilophyta; Polypodiopsida ili Polypodiophyta) uzgajanih u Vrtu između 1895.
i 2021. godine. Iako se stajališta stručnjaka o sistematici, klasifikaciji, nomenklaturi i taksonomiji papratnjača i danas uvelike razlikuju, u ovu su skupinu biljaka već uobičajeno uvrštene i preslice te psiloti. Rezultati pokazuju da je tijekom 126 godina u Botaničkom vrtu uzgajano najmanje 376 divljih i
uzgojnih svojta paprati iz 29 porodica. Nažalost, prekid redovitog rada („COVID-19-lockdown“),
snažni potresi te vremenski ekstremi koji su pogodili Zagreb 2020., nastavljajući se u 2021., imali su za
posljedicu i osiromašenje naših zbirki. Danas uzgajamo 86 svojti iz 20 porodica papratnjača.
Ključne riječi: Botanički vrt PMF-a u Zagrebu, povijesne zbirke biljaka, papratnjače, paprati,
preslice, psiloti
FOREWORD
In 2015 we launched the “Plethora of Plants” -series for Natura Croatica museum
journal, studying the trends in various increased or declining Botanical Garden plant
collections over the years. After my first thorough inventory of the temperate glasshouse plants (‘Aroideae’) in 2015, attempts were made to improve the glasshouse col-
134 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
lections on the basis of these surveys, which deduced the plant-lists as initially established, and identified the plant taxa that should be added to our recent collections, thus
“filling the gaps” in the inventories of families and/or genera. Statutorily protected as
a monument of nature and culture of the Republic of Croatia, and the City of Zagreb,
our Botanical Garden is dedicated to the preservation of the national botanical history.
It is thus important to try to replace the “missing parts of the puzzles” from our collections, bringing them closer to the original framework, as depicted in the earliest
reviews (Ettinger, 1892; Heinz, 1895-96).
INTRODUCTION
The comprehensive investigation of the plant collections in the Botanical Garden of
the Faculty of Science, University of Zagreb (below: “Botanical Garden” or “the Garden”) is continuing with inventories of indigenous, wild and cultivated ferns (Monilophyta; Polypodiopsida – Polypodiophyta), following the general principles established
in the first part of this series (Kovačić, 2015).
Ferns (Monilophyta; Pteridophyta – Polypodiopsida/Pteridopsida)
The classification of ferns sensu lato is considered very difficult, as briefly elaborated
in Kovačić (2015), just before the much-anticipated report of The Pteridophyte Phylogeny Group (PPG-1) was published in 2016. Alas, instead of bringing much needed consensus and order to the older group of “Filices”, the solutions proposed by PPG-1
provoked more disputes and (dare we say) new confusion to the World of Ferns.
In Kovačić (2015) I was dealing with combined historical data on our temperate glasshouse collections, ferns included (according to the to then-valid plant-bases The Plant
List, TROPICOS and IPNI), which I will not repeat here. But, it should be admitted, I was
truly astonished with the assortment of (not only) ferns in our original collections: according to Heinz (1895-96; pg. 14-15), “many species” from 21 fern genera and 6 families
(in my estimation, at least 35 taxa) of that time grew in our small Exhibition Glasshouse
back then, while in 2015 we had 25 taxa from 7 families (data in Kovačić, 2015). Investigation of our old paper-card database, established around 1952, revealed that after the
Second World War (WWII) we had a total of 13 glasshouse fern families, 26 genera and
53 species archived (Tab. 4 in Kovačić, 2015), out of which 7 fern families and 10 genera
were not present in 2015. However, these numbers are highly dependent on the nomenclatural authority used, which was for ferns always questionable. From 2015 until 2021
our glasshouse fern collection increased significantly, though the nomenclature problems
remained: consulting more than a single botanical authority brings linear growth to the
nomenclatural nightmare among ferns. Virtually, there are not two databases capable of
treating even our small fern collection in the same way: some species have more than 50
synonyms through several families, resulting in immense confusion.
As Professor Peter F. Stevens reminds us in the Angiosperm Phylogeny Website (APweb), terms such as higher and lower plants, or primitive and advanced characters, are
outdated. Nowadays, when talking about individual characters, terms such as plesiomorphic vs. apomorphic, or derived, are more suitable. All “land plants” (Embryopsida)
belong to a clade embedded in a predominantly aquatic, paraphyletic group of the
“green algae”: together, they construct the “green plants” (Viridiplantae) of our planet, in the broadest sense. No scientific consensus on the relationships among all aspects
Nat. Croat. Vol. 31(1), 2022
135
of plant life on Earth has been achieved. Accordingly, the relationships between the
“true” and “weird” ferns (actually, sophistically elaborated evolutional lineages, such
as Lycopodium, Selaginella, Psilotum or Equisetum) are still being much disputed. For
example, though morphologically astonishingly similar, whisk ferns (Psilotum spp.)
really are not descendants from the Palaeozoic Rhyniophytes – moreover, they are
entirely unrelated, and not nearly as old as previously thought. Whisk ferns are truly
ferns, as already accepted in the time of our Garden collections establishment.
Ferns in the Garden collections of the late 19th century
Though never citing the authority according to which he planned the layout of our
Garden, it is obvious that in imagining, planning, drawing, constructing and planting
the Royal Botanical Garden in Zagreb (as it was originally christened at its foundation,
during the Habsburg Monarchy), Professor Heinz (1861–1919), aided by his Chefgärtner
Vitĕslav Durchánek (1857–1924), followed Engler’s System. That, one of the prime
Darwinian phylogenetic systems of plant taxonomy based on evolutionary trends, was
developed by the famous German botanist, Der Altmeister Heinrich Gustav Adolf Engler (1844–1930), Heinz’s contemporary. Engler’s comprehensive system of Plant Life
classification with Orders and Families was commonly accepted, and it is still appreciated today. Widely called Syllabus der Pflanzenfamilien (or Syllabus of Plant Names;
Engler, 1892), it was the first “post-Darwinian”, at the time modern and complete,
revision of plant families to generic level in a single short publication. It was followed
by the monumental work Die natürlichen Pflanzenfamilien (The natural plant families;
Engler & Prantl, 1887–), gradually published during the years and containing much
more detail.
Though Heinz never mentioned Engler in his Guide, he states, as it goes without
saying (a fine German word is selbstverständlich): “Today we divide the Plants on Earth
into four large natural divisions, namely: 1. Myxothallophyta, 2. Euthalophyta, 3. Embryophyta zoidiogama, 4. Embryophyta siphonogama.” (Heinz, 1895-96, pg. 13). As those
terms are today abandoned, a short explanation might be in order: Myxothallophyta (or
Mycetozoa) was a polyphyletic grouping of so called “slime moulds”, evolution lineages of organisms that are, actually, unrelated, and today included mostly in the
Phylum Amoebozoa of the Eucaryotes. Euthalophyta (Thallophyta or Thallobionta) included fungi, lichens and algae, and – according to various views – even bryophytes,
bacteria and slime moulds. Thallophyta with “hidden” reproductive system was, together with ferns, incorporated into the “Cryptogamae”, as opposed to “Phanerogamae” – plants with “visible” reproductive system. Also, “Thallophyta”, with their
“undifferentiated (thalloid) bodies”, were opposed to “Cormophyta” – plants with
roots and stems. Embryophyta zoidiogama was Engler’s grouping (division) of mosses
(with hornworts and liverworts) and ferns (with equisetums and lycophytes). Embryophyta siphonogama (or Phanerogamae) consisted of gymnosperms (conifers, cycads,
Ginkgo, and gnetophytes) and angiosperms (all other “flowering plants”). This is the
Engler’s, Darwinian system, in its core (Buggs, 2021).
The second subdivision of Engler’s Embryophyta zoidiogama is the point where Heinz
starts his systematic description of the Garden collections: supraorder Pteridophyta,
with three orders – Filicales, Equisetales and Lycopodiales. The Order of Ferns (Filicales)
of the time comprehends two Suborders: “true ferns” (Filices) and “water ferns” (Hydropterides). From the first Suborder, various families were cultivated in the Garden.
136 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
Besides some 20-odd fern taxa (some only depicted with “sp.”) connected to the glasshouse cultivation from the Polypodiaceae family (published in Kovačić, 2015), Heinz
(1895-96) enumerates “many species” of Aspidium (val. Tectaria Cav.; but also of Croatian
flora, under other genera), Asplenium (incl. then separated Scolopendrium, of which Heinz
writes himself in 1892), Athyrium, Ceterach, Blechnum, Polypodium, Pteris “and others”,
living outdoors (“beside the grotto”; Photo-tab. 1), without a single species named. However, as we still today grow some “common” Croatian taxa at the same place (though
the “grotto” was removed between 1963 and 1965, when the rocks and boulders were
used for constructing the Sub-Mediterranean Rockery), it could be assumed that those
were similar. Table 1 (in on-line version of this text) is amended with Heinz’s data for
the ferns living outdoors in 1895, which were not published in Kovačić (2015).
b
a
c
d
e
Photo-tab. 1: Fragments of the Garden times long gone: a) The Grotto with indigenous ferns and
horsetails, around 1895; b) Gentleman observing young giant redwood (Sequoiadendron giganteum) with
some ferns beneath, before WWI; c) Warm glasshouse with Platycerium and other tropical ferns, after
WWI; d) Indigenous ferns in the Karstic rockery, after WWII; e) Botanical Garden ferns among other
indigenous plants presented at Zagreb International Garden and Flower Exhibition (today FloraArt), 1969.
Ferns in the Garden collections between 1900 and 1950
During the thorough digitalization of the Botanical Garden historic archives, financed by the Croatian Ministry of Culture in 2020, we recovered several forgotten
notebooks, sketchbooks and lists from the early 20th century (Photo-tab. 2), with collection registries of the time – among them, also some inventories of ferns. These
hand-written lists, filled with antiquated synonymy, are invaluable documentation of
our history, partially filling the 50-year gap of our plant inventories between 1897 and
1952 (explained in Kovačić, 2015). Some of the registries name certain ferns for the first
time in our Garden collections, and I have added them to this survey (Tab. 1). We also
recovered some lists of fern spores: 60 species ordered from unidentified sources in
1925/26, and 22 ordered in 1948 via Indici Semini of Delft, Prague, Oslo, Dijon, etc. Ferns
from these lists were not incorporated in Table 1, as we do not know their fate. There
are also several lists, mostly undated, ordering plant-labels for the collections, naming
some ferns among other things.
Nat. Croat. Vol. 31(1), 2022
137
a
c
b
d
e
h
f
g
Photo-tab. 2: a) Plant inventories written in the early 20th century, listing taxa after taxa once existing
in our collections. Many such notebooks are lost, while some of the retrieved carry only a part of the
families with taxa inventoried in some year, and are marked as “No. 2” or “B” – indicating that once
there were (at least) “1” and “A”. b) Fern registry, around 1900. cd) Perennials registry from 1904 with
the list of ferns. e) Registry of glasshouses from 191? (year illegible). f) List of 60 fern species spores “…
received in 1925, to be sown in January of 1926”, without their further fate known. g) List of warm glasshouse ferns from 1932. h) List of plant-labels for the glasshouse ferns, early 1930-ies.
138 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
As nothing of our ferns was officially peer-reviewed and published during the first
half of the 20th century, and all data on our collections remained in – today seriously
damaged – hand-written inventories, we do not know more about the listed plants:
whether they were purchased, received as gifts, grown vegetatively, or from spores;
their origin or their age. Also, complex classification of ferns was unknown to our old
records: in the hand-written inventories from the first half of 20th century, sometimes
very difficult to decipher due to the ornate handwriting and yellowing pages, all ferns
were listed among “Filices” (without families), and later under “Polypodiaceae”, until
the new paper-card database was established in the early 1950-ies. It is also challenging to follow the nomenclature of the time. For the inventories written before WWI it
is sometimes very difficult to found a valid name, while the author of taxon is missing.
Cultons could sometimes be found in old (predominantly German) horticultural books
and magazines, some today available on-line (for example, Species Filicum by Hooker,
1785-1865; Lexikon der Gärtnerei by Dietrich, 1837; Ferns and fern culture by Birkenhead,
1892). We do not know practically anything of the literature used by our predecessors,
who might have had their private books they took with them when they retired: after
WWII very little was left. We can also presume that they used the “bare” plant-names
as they found them in the Indici Semini of the time, as synonymy in the remaining
hand-written inventories is vast.
Ferns in the first Garden paper-card database (ca 1952 to 2000)
The list of the temperate greenhouse species (incl. ferns) published in Kovačić, 2015
(Tab. 2, p. S8) based on our old paper-card database (Fig. 6ab, pg. 376) was found to be
incomplete, not just because of the aforementioned hand-written registries found
meanwhile. As mentioned before, all paper-cards were in 2020 subjected to thorough
digitalization, which is explained in Budisavljević & Kovačić (2020). This work revealed many “hidden” entries (Photo-tab. 3). Besides amending the previously published data on glasshouse ferns (Kovačić, 2015) with data of the outdoor ferns, in Table
1 I also added those “hidden” names from the reused paper-cards, whenever it was
possible to reconstruct the original, erased name. However, we can be certain that
many such reused cards were thrown away during the course of time; some we were
unable to reconstruct, and some almost certainly went under the radar, so this historic review again could not be seen as complete.
Undoubtedly, the launch of the typed paper-card database initiated by Dr Ungar
in the early 1950-ies was a significant practical improvement in comparison to the
alphabetized hand-written inventories irregularly conducted before. However, during
the decades, this database became drawn in rejected, unused and reused cards, many
discarded as duplicates – overwhelmed with synonymy, many species inventoried
separately under several synonyms, enhancing the confusion. When I became employed in the Garden (September 1st, 2000), this old database was already divided in
to the “active” and “passive” parts (explained in Kovačić, 2015), which – I am sorry to
say – I helped, during the years, to entangle even more. In 2002 our students benevolently started to digitalize the paper-cards into the Flora Croatica Database (some, some
not), which led to new database separations and divisions, worsening the confusion
furthermore. Completing the thorough digitalization process in 2021 to produce a
brand new Online-Garden Database of his own invention, my colleague Alan Budis-
Nat. Croat. Vol. 31(1), 2022
139
a
b
c
d
Photo-tab. 3. Oldest fern names in the paper-card database: ab) Cystopteris fragilis (“Alps, 1949”) redetermination-card was found before the original data-card, overwritten under one of the reconstructed
entries, making this name the oldest recorded fern found in our paper-card database; c) Asplenium
viviparum (1949) and d) Asplenium ceterach (orig. Ceterach officinarum var. crenatum; 1950) follow.
avljević called the whole inventory system “The Quicksand Effect” (see Budisavljević
& Kovačić, 2020).
Ferns in the Garden collections of the modern days
Until the early 21st century, almost all classifications were heavily based on plant
morphology; as such, they differ from classifications based on molecular results. Furthermore, the use of morphology in combination with molecular data affects the relationships detected – in other words, one should use one (morphology) or another (molecular), not combining both (Stevens, 2021). The “clash” between the traditionalists
(morphologists) and the modernists (molecular analysts) is inevitable.
As we, the botanic garden lot, are working among living plants in our collections on
a daily basis, while the laboratory-based molecular lot often has very little contact with
the “real plants out there”, the criticism of the “molecular results” from “our side” is
often pointed in that way. Having been a member of the “garden lot” for almost 25
years now, I personally think that the latest results of this extensive and immensely
thorough research must be acknowledged, though, the LIVING PLANT can never be
less important than a RESULT of a method (even a molecular method). The amount of
140 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
scientific results published daily on any subject today is gigantic, and so would be the
literature citations on this matter, and on ferns in any sense, exceeding the length and
purpose of this text. To “bridge” this problem, I decided to refer here to the Angiosperm
Phylogeny Website (APweb), hosted by the Missouri Botanical Garden (USA) and maintained by Professor Peter F. Stevens, where an incredible amount of literature is ready
to found. The APweb is dedicated to research on (first of all, angiosperm) plant phylogeny and taxonomy, where all the latest references are regularly updated and presented in a way that could be understandable not only by the professional botanists.
Professor Stevens edits the ferns in APweb according to the results of The Pteridophyte
Phylogeny Group (PPG-1, 2016), with regular modifications to incorporate new results
and different views. Besides, in my opinion, his humorous and relaxed approach to
the perplexing nomenclatural entanglements, which will never be completely solved,
no matter what we do, is quite infectious (these days, in a good way). Clarifying that,
it is needless to say that the recent classifications are much more elaborate and much
less straight-forward than Engler’s, in Heinz’s times.
The modern clade of ferns and their relatives – [monilophytes + lignophytes] –
stemming out of molecular investigations is sometimes called the Euphyllophyta; those
plants share a morphologically common leaf pattern, called megaphylle or euphylle,
which varies considerably. Phylogenetically, ferns and their relatives – the monilophytes and lignophytes – are both well supported clades. According to the latest comparative data (details in APweb, 2021), the monilophytes (ferns) derived from the lignophytes (extinct progymnosperms and all extant woody plants, which gave rise to the
seed plants of today) between 482 and 385 million years ago. Until today, ferns and
gymnosperms divided greatly, having very little in common (e.g., cyanogenic glucosides, unlike any other plants out of their lineages).
Needless to say, this modern Clade of Ferns – Monilophyta – is not universally
accepted. It carries many former names (some pro parte), out of which APweb (2021)
gives full synonymy only to the Class Polypodiopsida Cronquist, Takht. & W.Zimm.
Acc. to the recent views, monilophytes include 47 families and a large group (clade) of
eupolypods I and eupolypods II, which includes most ferns. Members of this clade
changed their affiliations to one family or another more frequently than any other – a
fact that botanical garden employees, kept busy changing plant name-plates for visitors, know all too well (Photo-tab. 4).
Morphologically (but very much supported molecularly), up to 98% of all living
ferns are leptosporangiate: their sporangia arise from a single epidermal cell. The remaining 2% of species are distributed in no fewer than three further ‘major groups’ of
ferns: the eusporangiate or marattioid ferns (Marattiidae, Marattiaceae) with sporangia
rising from a group of cells, the horsetails (Equisetiidae, Equisetaceae), and the whisk
ferns (Psilotaceae and Ophioglossaceae).
Within the leptosporangiates, well over 70% of ferns are so called eupolypods (an
older term would be Order Polypodiales Link; as in very broadly defined family Polypodiaceae of the Christenhusz & Chase (2014) classification). It is interesting to emphasize that about one third of the eupolypods are epiphytes, ca 10% of all vascular
epiphytes. Though people commonly connect ferns with shady and moist habitats,
globally, desiccation tolerance is quite common in both epiphytic and terrestrial fern
sporophytes: their peltate scales may play a central role in the uptake of water – rather
as in Bromeliaceae. It is also found that 38-43% of all ferns, especially members of the
Nat. Croat. Vol. 31(1), 2022
141
[Marattiidae + Polypodiidae] clade, accumulate aluminium, and in a far higher percentage than any seed plants. Ferns also commonly have phytoecdysones, chemicals
they synthesize for defence again insects, also successfully shielding them against many
other herbivores.
Ferns are noted for their high incidence of polyploidy, involved in almost 31% of all
speciation events. Although polyploidy is common, the amount of DNA per chromosome tends to be conserved, unlike in angiosperms. Particularly large genomes occur
in Psilotales and some Ophioglossales, but in a few polypods.
a
b
c
d
Photo-tab. 4. Complex fern nomenclature and affiliations to families are seen in our Garden paper-card
database: some taxa, even Croatian native, changed names and families several times: a) Three name
changes of Polypodium cambricum. b) Four times re-written species and three times family of Dryopteris
villarii. Some names are unknown, left without a family: c) “Mycrosoryum pteropus” / “Microsorum
pteropos”. d) Many cards carry two valid names as synonyms: f.e. Pteris vittata = P. longifolia.
Review of the recent phylogenetic affiliations of ferns, with remarks on the
representatives in our Garden collections
The first phylogeny-based reclassification of ferns was proposed by Smith et al.
(2006, 2008), which has by now grown in detail (and changed) immensely. It is worth
mentioning that among the “ultra-modern” molecular fern phylogenetic classifications
there are different views (e.g., Christenhusz & Chase, 2014), as well as elaborated
disputes (Christenhusz & Chase, 2018; Schuettpelz et al., 2018) among specialists,
which Stevens (APweb, 2021) calls “splitting vs lumping”. Though we, the botanic
garden lot, possibly prefer the morphological over the molecular “way” in the systematization of plants (especially when it leads to frequent changes of names!), we are still
obliged to follow the recent literature and trends in research, to find a “least painful”
mid-way in bringing the topic closer to the widest audiences, while promoting and
popularizing botany. In an attempt to bring the “classic and modern” together, I should
try to follow Heinz’s lead from our first Garden Guide (descriptions of ferns in our
142 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
Garden collections, written in 1885-96) modified according to the latest views and
discoveries. In Table 1 I compared the recent views and affiliations on ferns comparing
three large plant-bases: APweb (2021, following PPG-1, 2016), the World Flora Online
(WFO; follow-up of The Plant List), and Plants of the World Online (PoWO), maintained
in the Kew Botanic Garden. For Croatian native species, I consulted Flora Croatica Database (FCD), to which our national legislation is connected.
a
b
c
d
Photo-tab. 5. True relatives: a) Adder's-tongue fern (Ophioglossum vulgatum), b) young whisk fern (Psilotum nudum), c) great horsetail (Equisetum telmateia) and d) ladder brake (Pteris vittata) in our Garden
collections.
“Weird ferns”
From the beginnings of “modern” plant classification, Psilotum and Equisetum had
been thought to represent lineages independent of each other, and unrelated to ferns.
Psilotum and its relatives are considered to be the most primitive living vascular plants
(APweb, 2021, with extensive literature). That is not the case, in spite of their “weird”
morphology (Photo-tab. 5).
EQUISETOPSIDA – EQUISETIDAE Warm.
EQUISETALES DC. ex Bercht. & J. Presl. – Equisetaceae Rich. ex DC.
As the circumscription of the fern clade has only recently become clear (though disputed), there is still no consensus on the position of Equisetum. There are three (molecular phylogenetic) hypotheses on the position of Equisetum among ferns, and many
details on their relationships are still waiting for explanations (APweb, 2021 with extensive literature within). Most supported seem to be the view treating the genus (family
and order) as sister to all other ferns (monilophytes), though there is also much support
for the theory suggesting the [Psilotales + Equisetales] clade.
There is a long fossil trail of Equisetum-like plants (“equisetophytes”) from the Upper
Devonian onwards, while “true” Equisetum is known from the Triassic. Fascinatingly
little change has been undergone by the horsetails in their basic morphology over a very
long course of time, visible even in our little fossil collection (Photo-tab. 6 in Suppl.);
Equisetum is considered to be one of the most successful genera of extant vascular plants
of all times. Most of the apomorphies of the genus are to be seen in the context of variation in the taxa of extinct equisetaceous Calamitaceae, rather than other extant ferns.
The clade containing Equisetum (some suggest the existence of a [Equisetidae + Ophio-
Nat. Croat. Vol. 31(1), 2022
143
glossidae] clade) has probably been separated from the other monilophytes since the
Permian, ca 250+ million years ago. However, the modern species of Equisetum seem to
have separated as late as the Caenozoic, before ca 65 million years. The single extant
family has a single extant genus (Christenhusz et al., 2019).
Among the many traits unique to horsetails, or connecting them with other ferns,
are: having both erect and creeping stems without secondary thickening; non-mycorrhizal roots; accumulation of SiO2; leaves not photosynthetic, whorled and small, simple
and 1-veined; spores with circular aperture; n = 108. It is interesting to emphasize that
the leaves of Equisetum – so different from a fern’s ‘fronds’ – may be secondarily simple.
Early members of this clade had secondary thickening, while their spores were very
different from those of today (3-ridged in Calamites, unridged in Equisetum). Aided by
their elaters, spores of Equisetum can “jump” up to 1 cm in the air as they dry, or “walk”
by short random movements along the ground.
It is worth mentioning here that the stomata of a few monilophytes, including Equisetum (and perhaps also Psilotum), do not respond to red light; they are likely to be immobile (permanently closed), especially when older, because of the rigid radiating riblike silica thickenings of the subsidiary cells. In total absence of silicic acid, stomata can
develop normally, as well as the whole plant.
In their distribution, horsetails are nearly cosmopolitan: there are 18 species in total,
out of which we in Croatia have 9 (FCD, 2021).
Heinz (1895-96) described “several species of Equisetum growing by the grotto” in the
Garden, without further details (Tab. 1). Older inventories rarely mention this genus.
The first entry in our recent database is from 1961, of a freshwater E. fluviatile L. (already
noted in the inventory of 1958), which since then has vigorously filled one of the ponds
of our systematic field (Photo-tab. 7 in Suppl.). Equisetum arvense L, common throughout
the temperate Northern Hemisphere, was never inventoried, as it grows wild along the
Garden’s disturbed places. Recently, we are trying to keep it in our fernarium collection,
primarily for educational purposes. Surprisingly, even E. telmateia Ehrh. does not live
long with us in recent decades, since the summers become hotter and dryer. Short-lived
in our fernarium were E. sylvaticum L. and E. hyemale L. (vulnerable (VU) and strictly
protected in Croatia), which are occasionally brought from wild localities. According to
Dr. Regula (pers. comm.), other native horsetails were also brought from field research
occasionally and placed in the cold greenhouse, but they never survived a single year.
In order to show more horsetails to our visitors, we recently purchased E. giganteum ‘El
Tabacal’: it proved to be vigorous, growing happily with the rest of the ferns in our fernarium, though it should be monitored and kept from spreading too much.
Horsetails tend to grow in ecologically rather stressful habitats. They have so-called
convective ventilation from the stems into the rhizomes, allowing the “equisetaceous”
plants to penetrate deeply into the anoxic substrates, commonly favoured by this group.
However, species like E. hyemale lack that interconnected air space and have no convective ventilation, yet still can grow in anoxic, partly submerged conditions.
PSILOTALES & OPHIOGLOSSALES (in different versions)
The close affiliation of these plants with ferns, today commonly accepted, was indeed surprising, though some older authors had earlier compared Psilotum with some
extinct and extant ferns (comprehensive literature in APweb, 2021).
144 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
The clade consisting of [Psilotum + Ophioglossum] is perhaps sister to all other ferns.
Ophioglossidae could also be comprehended as an equivalent to the class Psilotopsida,
or one of the four subclasses of Polypodiopsida (ferns), containing the orders Psilotales
and Ophioglossales. This relationship was only confirmed by molecular phylogenetic studies (APweb, 2021), and it is morphologically highly disputed. Like Equisetum,
both Psilotum and Ophioglossum have erect and creeping stems, but their gametophytes
are subterranean, axial and non-photosynthetic (mycorrhizal). Both have large genomes which are similar in terms of apomorphies to those of seed plants, and share
mycorrhizal associations with the fungi of glomalean Glomus group A. The age of this
clade (if such exists, as APweb (2021) puts it) is estimated to be ca 275 million years.
PSILOTALES Prantl – Psilotaceae J.W.Griff. & Henfr.
If taken as such, this Order contains a single family: Psilotaceae, with two genera.
These are epiphytes without true roots; leaves are small and simple (laterally flattened
in Tmesipteris and highly reduced in Psilotum); spores are kidney-shaped and monolete.
The “whisk ferns” are of pantropical distribution; their age is estimated at between 80
and 72 million years (APweb, 2021, literature within).
The genus Tmesipteris is notoriously difficult in cultivation with very specific requirements, and we never had it growing in our Garden. Its 15-odd species (depending
on the authority) are delicate, low-height epiphytes growing on the trunks of tree-ferns
(Dicksonia, Cibotium) and occasionally on shady, moist rock-faces. The most recognizable species, Tmesipteris obliqua Chinnock, is famous for its genome gigantism (Hidalgo et al., 2017): the largest known of all ferns.
The genus Psilotum, which contains two species, has also never been inventoried in
our collections since Heinz’s times (1895-96): there is just a single note of spores received in 1925 (Tab. 1). Preparing the exhibition on the evolution of plants in 2018
(Kovačić, 2018) in our Garden pavilion, we constructed a “Devonian Diorama” (Photo-tab. 8 in Suppl.) along with Carboniferous and Cretaceous, designed in accordance
to the data and drawings of the famous Lower Devonian Rhynie Chert in Scotland.
As our gardeners were making some simple Rhynia-models for this Diorama, we came
to the conclusion that the psilotums would be ideal to show to our visitors (mostly
children) some similar extant plants, not related to the extinct Devonian rhynias (Psilophytales). Through the European Botanic Garden Consortium (EBGC) network we asked
for help, which came immediately from our neighbours, the Budapest Botanical Garden of the Eötvös Loránd University (ELTE). Our colleagues sent us several young
Psilotum nudiflorum (L.) P. Beauv. plants, which we incorporated into our exhibition,
and afterwards into our glasshouse collection.
OPHIOGLOSSALES Link – Ophioglossaceae Martinov
This order and family of the peculiar-looking “moonworts” consists of 4-10 genera
and 55-125 species (depending on the authority), while their classification is still on-going (APweb, 2021). Most of these species have fleshy roots and a single, simple leaf
blade with netted venation; spores are short-lived. They are nearly worldwide in their
distribution, with age estimated at between 256 and 162 million years (APweb, 2021;
literature within).
The eponymous genus Ophioglossum holds at least two records in the plant kingdom:
for the world’s smallest land fern (O. malviae Patel & Reddy: ca 1 cm) and the largest
Nat. Croat. Vol. 31(1), 2022
145
known number of chromosomes (O. reticulatum L.: n = 720). Years could pass before
the plant sends a leaf above ground, with (or without) a spore stalk. The non-photosynthetic gametophyte meanwhile lives underground, sustained by peculiar soil fungi, not known from any other plants.
In the Croatian flora we have two genera of this family, living shortly growing in the
Garden (perhaps not, but when the plants withdraw underground it is quite difficult to
maintain the “empty pot” (which our gardeners soon forget), so they were regularly
“presumed dead”). Ophioglossum is represented by three species, out of which O. lusitanicum L. is critically endangered (CR) and strictly protected (FCD, 2021). Most common
of three Croatian species in this genus, Ophioglossum vulgatum L. (Photo-tab. 5 in Suppl.)
is sporadically brought to the Garden from the field, where it can be found in its “regular places” – as is Botrychium lunaria (L.) Sw. The genus Botrychium is presented in our
flora with two species, out of which B. matricariifolium (Döll) A. Braun ex W.D.J. Koch is
critically endangered (CR) and strictly protected (FCD, 2021). The leaves of Botrychium
can take up to five years to develop, perhaps a record for land plants (APweb, 2021).
“True” ferns
The position of the first “true” ferns, Marattiidae/Marattiales, is still disputed. Some
molecular phylogeny research suggests that the [Marattiales + Psilotales] clade could
be sister to all leptosporangiate ferns. However, the [Marattiidae + Polypodiidae] clade
is more widely accepted. Ferns of this group share compound leaves (fronds) with
scales; the gametophyte is green (photosynthetic) and above-ground; plants very often
accumulate aluminium. Various authors have suggested ages of this clade between 337
and 329 million years (APweb, 2021 and literature within).
MARATTIIDAE Klinge – MARATTIALES Link – Marattiaceae Kaulf.
Eusporangiate Marattiaceae are considered to be one of the most “primitive” living
families of ferns, diverging from other ferns very early in their evolutionary history
and quite different from other ferns common in temperate parts of the world. Ferns of
this lineage are richly represented in the Permo-Carboniferous fossil records, about 300
million years ago (Photo-tab. 9 in Suppl.). At least five marattiaceous clades lived from
side to side of Pangaea before its splitting (APweb, 2021).
Today, the Order consists of just one family with 6 genera and 110 species of pantropical distribution, the estimated age of which is between 236 and 201 million years.
Many have massive, fleshy rootstocks and largest known fronds of any (except tree)
ferns: up to 9 meters long in Angiopteris javanica C.Presl.
Heinz (1895-96) states “Angiopteris longifolia” (val. A. evecta (G. Forst.) Hoffm.) to
have been living in the glasshouse of the time, but there is no trace of this family’s
representatives afterwards. In 1925, the spores of several species from Angiopteris and
Marattia (incl. Eupodium and Ptisana) were inventoried as having arrived, but nothing
further. In 2016, we tried to grow A. evecta from spores, with poor success.
POLYPODIIDAE Cronquist, Takht. & Zimmer.
The largest clade of extant ferns on Earth, the leptosporangiate Polypodiidae have
very strong support and share many common characteristics (APweb, 2021). The polypodiidian gametophyte is cordate and above-ground, with more or less exposed an-
146 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
theridia. Blue light stomatal opening response is absent in this group, unlike in all
previously described lineages. Sporangia bear 64-800 spores. Tolerance of extreme
desiccation is sometimes facultative and scattered through this clade, which occurs in
gametophytes, too. The estimated age of Polypodiidae varies from 350 to 170 million
of years, according to different reconstructions.
OSMUNDALES Link – Osmundaceae Martinov
This lineage seems to have special position within the leptosporangiate ferns, as
their sporangia bear some eusporangiate features, and are strongly supported as being
sisters to the rest. A large portion of Osmundales became extinct at the end of the
Permian, especially of genera that lived in various damp habitats. For such a small
group, there is an extensive literature depicting complex classification relations of
extant and extinct osmundaceous taxa (APweb, 2021 and literature within).
Today, there is a single extant family with 1(-6?) genera and 18 species of nearly
cosmopolitan distribution. Some of the mutual features are accumulation of SiO2 ,
green spores and n = 22. The age of this fairly isolated clade is estimated to between
265 and 200 million years, but it has also been suggested that the Osmunda clade in
particular originated as early as the Carboniferous (ca 323 to 305 million of years). The
extinct osmundaceous taxa are particularly interesting, while their fossils have been
extremely well preserved: for example, the fossil from Sweden, dated to ca 180 million
of years, has anatomy that remarkably resembles living Osmunda (“Claytosmunda“)
claytoniana L.
A critically endangered (CR) and strictly protected member of the Croatian flora
(FCD, 2021), Osmunda regalis L. is the largest and most impressive European fern, a
deciduous species bearing separate dimorphic (fertile and sterile) fronds. Results of
fossil examination show that neither ploidization events nor notable amounts of gene
loss have occurred in the genome of the “royal fern” since the Early Jurassic, ca 180
million years ago – which is astonishing to our Garden visitors of all ages, observing
the living plants in the Garden since Heinz’s times (1895-96). The oldest living in our
collections today was brought in 1962 from the Borlin locality near Karlovac, where it
vanished since then, due to the unwise habitat changes. Later samples were brought
from the Croatian Banovina region, always as living plants (Photo-tab. 10 in Suppl.).
Of the mature plant we brought in 2016, I took some spores in 2017 and grew several
“babies” in our laboratory, which we planted in the spring of 2021 around their “mother”. Heinz (1895-96) also cites “Todea sp.” of this family, as living in the greenhouse.
The later, hand-written inventories named several representatives (incl. O. gracilis =
val. O. spectabilis Willd. and O. cinnamomea = val. Osmundastrum cinnamomeum (L.)
C.Presl), living in the Garden of the time, as seen in Table 1. The list of spores received
in 1925 depicts three more: Osmunda barbara Moore (? Thunb.; val. Todea barbara (L.)
T.Moore) and O. claytoniana L., which was noted as “possible”, so I presume that it
sprouted, but the further inventories are lost.
The polypoid clade consisting of [Hymenophyllales, Gleicheniales [Schizaeales
[Salviniales [Cyatheales + Polypodiales]]]], is estimated to be between 286 and 273
million years old. The most important trait separating these ferns from the eusporangiate Marattiales (and “transitional” Osmundales) is that their stems have a single
apical cell: they are true leptosporangiate ferns.
Nat. Croat. Vol. 31(1), 2022
147
HYMENOPHYLLALES A.B. Frank - Hymenophyllaceae Martinov
This group of leptosporangiate ferns holds just one family, with 2(-34!) genera and
ca 435 species, which are most often epiphytes without mycorrhizal fungi, and filamentous or ribbon-like gametophyte. The age of this group is broadly estimated at
between 243 and 176 million years. Hymenophyllaceae are not exclusively tropical
plants, but are very vulnerable to desiccation which limits the range of habitats in which
they can survive. Often called the “filmy ferns”, they are restricted to humid or constantly sprayed places (for instance, under waterfalls or by springs).
Traditionally, two genera of this family were recognized, as in Heinz (1895-96): (1)
the mostly terrestrial Trichomanes s.l. (now a central genus of the clade Trichomanoideae) with tubular involucres, and (2) the epiphytic Hymenophyllum (a single genus of the
clade Hymenophylloideae) with bivalved involucres. Heinz (1895-96) cites representatives of both genera as „growing in the greenhouses” of the time: "Trichomanes
radicans" (val. T. speciosum Willd.) and Hymenophyllum tunbrigense (L.) Sm. The latter
is of worldwide but discontinuous distribution, in accordance with its special living
requirements. It was recognised above Rijeka (the Rječina) for Croatia by Schlosser &
Vukotinović (1869), but nobody since them has seen it at the alleged locality. Decades
later, it was found again (Hirc, 1903) in Samoborsko gorje, today a Nature Park. In
spite of several “sightings” during the 20th century, Trinajstić (1994) claimed that after
the 1930s nobody actually had a proof of finding this elusive fern in Croatia.
In our paper-card database there is no trace of this family members being grown in
the Garden. I must admit that it is a mystery to me where and how the “filmy ferns”
were kept in our Garden in the late 19th century, as we do not have the means to maintain them today, nor have they ever appeared in any of the Garden inventories after
Heinz’s in 1895.
SCHIZAEALES Schimper
Members of this order share dimorphic (fertile and sterile) fronds. There is a lack of
well-defined sori: schizeaceous sporangia have a horizontal annulus encircling the top
of the sporangium. The age of this group is estimated to between 218 and 184 million
years; it includes 3 morphologically well-distinguished (though disputed) families,
with 4 genera and up to ca 200 species. According to the older views, the order consisted of just a single family, the eponymous Schizeaceae s.l.
Ca 100 species of the monotypic Anemiaceae Link family are widespread, typically
terrestrial or epipetric ferns. In his times, Professor Heinz (1895-96) tagged “Aneimia
Phyllitidis” (val. Anemia phyllitidis (L.) Sw.) as living in the greenhouse, present also in
later inventories (Tab. 1). The monotypic family of climbing ferns, Lygodiaceae C.Presl,
is distributed along tropical and warm temperate regions. The rachis of each frond is
long and flexible, with indeterminate growth, forming climbing or trailing vines. The
family has only one genus, Lygodium, with ca 20 species, not mentioned by Heinz (189596). In the hand-written registries L. japonicum (Thunb.) Sw. was inventoried several
times (Tab. 1), as well as sprouting among spores received in 1925. Recent specimens,
grown in the Garden laboratory, are living in our tropical Victoria glasshouse (Photo-tab. 11 in Suppl.). We have never had representatives of the pantropical-temperate
family Schizaeaceae Kaulf. in the recent sense (two genera) in our collections.
148 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
SALVINIALES Link
This Order encompasses aquatic and semiaquatic ferns without mycorrhizal fungi.
They have dimorphic (fertile and sterile) fronds and sporocarps; the plants are heterosporous; sporangia lack annulus and contain a single megaspore per megasporangium.
The age of the order is estimated to be between 186 and 153 million years, including 2
families (Photo-tab. 11 in Suppl.) with 5 genera and 80-odd species.
Mostly tropical to warm temperate, Salviniaceae Martinov is a family of free-floating ferns, carrying sessile, 2-ranked, simple fronds with distinctive stomata. It holds
two genera, both of which we have representatives in our collections, always inventoried among “water plants”, never among ferns. The native Croatian species, Salvinia
natans (L.) All., has lived in our outdoor and indoor basins since Heinz’s (1895-96) time,
being required for teaching water repellence of fronds to our students. Though in
Croatia not endangered in the wild (even a serious nuisance in fishponds and fish
farming basins), S. natans is nevertheless statutorily strictly protected, due to the
EU-legislation (Bern Convention). During the years, we also had several exotic representatives of the genus Salvinia (Tab. 1), which are challenging to keep separated outdoors, due to the transfers between the basins. The American genus Azolla (“sp.”) was
also known to Heinz (1895-96): we always keep A. filiculoides Lam. in our collection.
All Azolla species have obligate association with Nostoc azollae cyanobacteria, important
nitrogen fixers (APweb, 2021). A second family, Marsileaceae Mirb., includes three
tropical-temperate genera of (semi)water ferns. Fronds are simple, linear or divided to
2/4 leaflets, characterized by bean-shaped sporocarps and heterosporangiate sori; long
and slender rhizomes creep along or beneath the ground. Out of the three genera, we
have two in the Croatian flora. The critically endangered (CR) and strictly protected
Pilularia minuta Durieu was never inventoried in our Garden database, as it never lived
long (Dr. Regula, pers. comm.). Endangered (EN) Marsilea quadrifolia L. inhabits our
outdoor basins since the time of Heinz, who also mentioned some unnamed “other
species”. We also grew several “water clovers” in our more recent collections (Tab. 1),
but M. quadrifolia is the most stable; one of 20-odd strictly protected “Natura 2000”-Croatian plant species the conservation of which is important for the EU. We acquired the
monotypic Regnellidium diphyllum Lindm., representative of the third Marsileaceae-genus, for the first time in 2021 from Bonn Botanical Garden (Germany).
CYATHEALES A.B. Frank
Depending on the authority (literature in APweb, 2021), the order of “tree-ferns”
has a single family, Cyatheaceae, with 13 genera and ca 745 species, while other views
recognize 8 families (optionally, subfamilies of Cyatheaceae), which are predominantly monotypic. The age of this order is estimated at between 187 and 109 million years.
Cyatheaceae Kaulf. in the modern sense – out of which seven smaller families were
extracted – is a family of pantropical ferns with large fronds. This family includes the
tallest extant ferns on Earth, some growing up to 20 m high. As suggested by PPG-1
(2016), three genera with more than 600 species are true members of Cyatheaceae s.s.:
Cyathea, Alsophila and Sphaeropteris, which representatives in the Garden are listed in
Table 1. The newly established family Cibotiaceae Korall is monotypic: its 10 species
are scattered through tropical Central America, Southeast Asia to Malesia and Hawaii.
Several species were inventoried in the Garden before WWII (Tab. 1). Members of
Dicksoniaceae Bower nom.cons. never accumulate aluminium; their long, tapering
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hairs are composed of cells arranged end to end, unlike the scales characteristic for
members of the Cyatheaceae s.s. family. Three genera (Calochlaena, Lophosoria and Dicksonia) with ca 40-50 species have recently been classified in this monophyletic family,
out of which we occasionally have dicksonias (Tab. 1; Photo-tab. 12 in Suppl.). The rich
fossil record originates from all continents of ex-Gondwana, while their recent distribution is much narrower: tropical America, St Helena, Malesia to the Antipodes and
New Caledonia. It seems that some extant Dicksoniaceae-species have estimated ages
older than the islands they currently inhabit, so understanding the biogeography of
the family is still to be solved (APweb, 2021). We have never had representatives from
the rest of the “tree-fern” families in our collections, or else they were very short-lived.
POLYPODIALES Link
According to PPG-1 (2016), but not universally accepted, this mega-order includes
25 clearly separated families, and two groups of families of which the mutual boundaries are (so far) highly disputed. Polypoid ferns comprise nearly 90 % of all extant
fern species.
A number of small, often monotypic families with ambiguous positions are described (and disputed), leaving the total number of (many!) genera and species highly
dependent on the authority consulted. According to APweb (2021, literature within)
the morphological traits common for the members of Polypodiales are very general:
they share, for example, black, wiry roots; dorsiventral rhizome; vertical annulus interrupted by stalk and stomium, and age estimated between 260 and 150 million of
years. As confusing and entangled as the systematic and classification of this large fern
group always was, it is clearly reflected in the chaotic paper-cards from our old Garden
database: names of species and affiliations to families were sometimes several times
changed, with various synonyms marked as non-valid. In Tab. 1 I have tried to affiliate
our past and present ferns of this large Order to the recent nomenclature, as much as
possible. WFO and PoWO disagree in affiliations of species to genera, and even families, which is truly confusing. Consequently, I have arranged the families alphabetically, according to APweb (2021), and genera according to PoWO, while WFO is soon to
be reassembled (as of December, 2021). Still, it is quite confusing in relation to our own
database, so I added the affiliation to family with each species, if it is different to APweb/PPG-1.
Most of the Garden fern inventories name polypoid taxa and families.
Out of the monotypic Lonchitidaceae Doweld family we had one species (Kovačić,
2015) out of two in total (possibly many more, according to the other sources): Lonchitis hirsuta L., affiliated to Pteridaceae and then to Dennstaedtiaceae Lotsy. The latter
family is much disputed, the range of genera changing many times, hard to follow even
in our humble Garden database. According to PPG-1 (2016), 10 genera with 265 species
belong to Dennstaedtiaceae, out of which the most common in our collection is Pteridium aquilinum (L.) Kuhn. This Croatian native fern was never attributed to this family
in our national Floras (or in the Garden), but first to Polypodiaceae, and then to Hypolepidaceae (FCD). To add taxa to Dennstaedtiaceae in our collection, in 2016 we grew
Hypolepis millefolium Hook. and Histiopteris incisa (Thunb.) J. Sm. from spores (Photo-tab. 13 in Suppl.).
Pteridaceae E.D.M.Kirchn. is a family of worldwide distribution (according to PPG1, 2016), which does not exist according to other authorities (for example, FCD: single
150 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
Croatian species of Adiantum, A. capillus-veneris L., is affiliated to Adiantaceae). Ferns
of this clade have bilateral spores; there is no indusium; gametophyte is ribbon-like,
and mycorrhizae are not present. They could be epiphytic, xeric or aquatic. Nine genera (disputably) belong here, out of which we have some growing wild in Croatia, and
many in the Garden collections since its foundation (Kovačić, 2015; listed in Tab. 1)
Today, 53 genera with 1210 species are included in Pteridaceae sensu PPG-1 (2016; via
APweb, 2021), with most gathered in Pteris (250), Adiantum (225 – in FCD Adiantaceae),
Cheilanthes (100 – in FCD Adiantaceae) and Pellaea (40).
Besides the taxa of this group which were published in Kovačić (2015), we grew
some new plants in 2016: for example, Llavea cordifolia Lag., Hemionitis palmata L., Cryptogramma acrostichoides R.Br., and a rare (or rarely found) species of Croatian flora,
Anogramma leptophylla (L.) Link (in FCD under Adiantaceae; depicted in Kovačić, 2017).
Alongside the data already published for the greenhouses, Heinz (1895-96) does not
list any particular species as growing in the open. In the hand-written records, many
species of this family could be recognized, but never classified as such, only under
other families. For example, the genus Gymnogramma Desv. should be a full synonym
of Hemionitis L., but the particular species often inventoried in our olden collections,
G. aurea (Willd.) Desv., belongs to two different genera: Cerosora according to PoWO,
and Pityrogramma according to WFO (Tab. 1)! Likewise, Pteris palmata Willd. belongs
to Hemionitis according to PoWO, and Doryopteris according to WFO. Gymnopteris repanda (Blume) Christ, inventoried in 1958, could be Bolbitis repanda (Blume) Schott of Polypodiaceae (PoWO), while WFO recognizes only G. repandra Christ – nevertheless, this
genus, Gymnopteris, is a synonym of either Leptochilus, Polypodiaceae (PoWO) or Hemionitis, Pteridaceae (WFO)… etc., etc… There are many more examples of this complex
systematic in the Polypodiales, all based on molecular results, not on morphology (to
be clear), as seen in Table 1 (some representatives of our collection are shown in Photo-tab. 14 in Suppl.).
Where the story of the well-defined (if disputed) families of Polypodiales ends,
another one starts: that of the large and unwieldy clade of Eupolypods [Eupolypod I
+ Eupolypod II] sensu PPG-1 (2016, via APweb, 2021). This heterogeneous group includes most of the extant fern taxa, scattered through many families (all heavily disputed). The “eupolypoid” fronds are to 1.5-times pinnate; spores are monolete and
reniform; x = 41. The age of this group has been estimated between 116 and 105 million
of years.
Eupolypod group I (‘Polypodiineae’) is a clade (suborder) which includes 9 families: [Didymochlaenaceae [Hypodematiaceae [[Nephrolepidaceae + Lomariopsidaceae]
[Dryopteridaceae [Tectariaceae [Oleandraceae [Davalliaceae + Polypodiaceae]]]]]]].
Members of this clade are quite commonly epiphytic, sharing many anatomical and
morphological characteristics (such as dense rhizome scales, perhaps protecting the
plants against desiccation and aiding in the absorption of water and nutrients). Many
taxa of this clade changed affiliations in our Garden database during the years – as
described in Kovačić (2015; pg. 369), shortly before the results of The Pterodophyte
Phylogeny Group (PPG-1) research were published in 2016.
In the past, our Garden “eupolypods-1” were by default affiliated to Polypodiaceae
J. Presl & C. Presl (sensu lato – that is, sensu recent “Polypodiineae”). Most of Heinz’s
(1895-96) outdoor ferns (Croatian native, and exotic) initially belonged here, many of
them having been transferred to other families, as literature authorities changed.
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Though Heinz recognized fern families of that time, eupolypoid species in our
hand-written inventories were (together with all other ferns, equisetums and lycopods)
for decades later inventoried under “Filices” (for convenience, I presume), which were
after the WWII renamed “Polypodiaceae”. In the “new” paper-card database, founded
in 1952 (with some back-notes to 1948) ferns were finally placed into proper families.
Beside the glasshouse species in our historic collections (Kovačić, 2015), Heinz
(1895-96, p. 15) listed under Polypodiaceae also other native ferns, growing at the time
“…around Grotto (Photo-tab. 1): Aspidium, Asplenium, Athyrium, Ceterach, Scolopendrium, Blechnum, Polypodium, Pteris, etc.” (Tab. 1). As he rarely affiliated species of ferns
to the genera, we cannot say today which were there. For example, “Aspidium filix
mas” is Dryopteris filix-mas for sure; “Ceterach” is almost certainly Asplenium ceterach;
while “Scolopendrium” refers to Asplenium scolopendrium of the "Phyllitis /sub/clade".
Many of these ferns today do not belong to the Polypodiaceae family sensu PPG-1
(2016), as seen in Table 1. However, many disagree (e.g.,PoWO).
So many Polypodiaceae members in the modern sense (reduced to “only” 63 genera)
suffer the enormous lists of synonyms and “accepted names” in various printed sources and online databases. The WFO accepts one, PoWO the other, while the Croatian
FCD – to which national legislation is related – often gives a third synonym of the same
taxon; not to mention that APweb sometimes does not agree with any. The first row of
Polypodiaceae family in Table 1 already illustrates this. The “crassifolium” polypoid
fern in our collection could belong to at least four genera: following Linnaeus’ Polypodium, to Pleopeltis (in our Garden database), Niphidium (according to WFO) or Campyloneurum (according to PoWO). The confusion over the changing names of just two of
the existing ferns in our temperate glasshouse illustrates this (Photo-tab. 15 in Suppl.).
Succeeding the recent views (APweb, 2021 following PPG-1, 2016; amended with many
since then published results of various fern investigations), I tried to revise some of our
confusing historical nomenclature and synonymy (Tab. 1), partially published in
Kovačić (2015).
Almost 90% of all Polypodiaceae are epiphytic (called the “grammitid” ferns), promoting this group as the major “hanging-on”- clade of the Monilophyta in general.
Though the first epiphytic fern coming to our minds is probably the giant “stag horn”
(Platycerium spp., indeed a member of this family), the majority are rather small, “twig”
epiphytes (APweb, 2021). Also, there is apparently the secondary association with
mycorrhizal ascomycetes, which has developed in the sporophytes (unlike in the epiphytic Hymenophyllaceae, which are non-mycorrhizal), and the polypoid epiphytic
ferns seem to be dependent on this association. Many of them trap litter, for example
Drynaria and Platycerium (examples in Photo-tab. 16 in Suppl.).
Cosmopolitan Dryopteridaceae Herter nom.cons. also are mostly epiphytic ferns,
with >3.5 times pinnate fronds. They include 26 genera and ca 2,135 species (PPG-1,
2016): the largest genus is Elaphoglossum (620-795 species, of which we have never, since
Heinz’s times, had representatives), followed by (also Croatian native) Polystichum
(500) and Dryopteris (400). Heinz (1895-96) had these ferns listed under the Polypodiaceae family of his time.
Glasshouse representatives – according to the systematic views before PPG-1 (2016)
– were published in Kovačić (2015). Species found in our recovered hand-written plant
registries, Croatian native and outdoor species, as well as newly grown and acquired
taxa, are listed in Table 1. Genus Cyrtomium is a good example of the mixed synonymy
152 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
in this family, of which the members often changed their affiliation: in our database,
at first, mostly because of the lack of literature – and then, lately, because of the plethora of sources. Three species which we grew in our collections were systematized
under three different families: first as Polypodiaceae, then Aspidiaceae, and finally
Dryopteridaceae. The ‘Japanese holly-fern’ (Cyrtomium falcatum (L.f.) C.Presl) is a newcomer to the Croatian flora, for the first time noted as an escapee from cultivation on
the walls of the Old City in the Island of Rab (Trinajstić & Španjol, 1994), and by now
scattered down the Croatian coast to the Old City walls of Dubrovnik (FCD, 2021).
Croatian common Dryopteridaceae members include Dryopteris with 10 species (examples in Photo-tab. 17 in Suppl.), and Polystichum with four species and two hybrids
(Photo-tab. 18 in Suppl.), one of which is a statutorily strictly protected endemic hybrid
P. x illyricum (Borbás) Hahne (Tab. 1), also popular in horticulture.
Monotypic “PPG-1-families”, excluded from Polypodiaceae s.l. on the basis of a
single genus (or even species), are disputed – also in our collections. For example, the
pantropical Didymochlaena truncatula (Sw.) J.Sm. (with 20-odd synonyms) gets its own,
monotypic family Didymochlaenaceae L.-B. Zhang & L. Zhang, which is not recognized
by either PoWO (Polypodiaceae) or WFO (Hypodematiaceae). Professor Heinz (189596) mentioned “Didimochlaena” under Polypodiaceae, and afterwards Didymochlaena
sinuata Desv. (val. D. truncatula (Sw.) J.Sm.) could be found just once, in the hand-written inventory of 1904 (Tab. 1). The predominantly South-East-Asian Davalliaceae M.R.
Schomb. are also monotypic (Davallia with 65 species) according to PPG-1 (2016), while
other views divide Davallia into up to seven genera, and both principles are equally
disputable (Christenhusz & Chase, 2014). Representatives of this family in our glasshouses were published in Kovačić (2015): since then, we grew two new species from
spores (Photo-tab. 19 in Suppl.).
PoWO places Tectariaceae Panigrahi to the large Polypodiaceae family. These (often
climbers) are pantropical, with 7 genera out of which the largest is Tectaria (210 species).
In addition to our old Tectaria cicutaria (Photo-tab. 19 in Suppl.), in 2016 we grew T.
zeylanica (Houtt.) Sledge (syn. Quercifilix zeylanica) from spores, a pretty little fern suitable for terrariums.
Family Nephrolepidaceae Pic.Serm. contains the sub/tropical genus Nephrolepis with
20 or 30-odd taxa, which PoWO again attributes to Polypodiaceae. In the Garden database affiliated to Davalliaceae or Dryopteridaceae, our glasshouse ‘sword ferns’ were
published in Kovačić (2015), while some new or previously “misplaced” plants (Photo-tab. 20 in Suppl.), are listed in Table 1. It is interesting that, among so many ferns of
his time, Heinz did not mention this horticulturally famous genus as existing in our
collections of 1895, which is hard to believe: there are at least a hundred different cultivars of ‘Boston fern’ (N. exaltata (L.) Schott) alone, many regularly inventoried in our
later handwritten registries (though the “names” of these cultons today are mostly
unknown and difficult to trace, Tab. 1).
Eupolypod II (‘Aspleniinae’) is a clade/suborder that includes 11 families: [Cystopteridaceae [[Rhachidosoraceae [Diplaziopsidaceae [Desmophlebiaceae [Hemidictyaceae +
Aspleniaceae]]]] [Thelypteridaceae [Woodsiaceae [Athyriaceae [Blechnaceae + Onocleaceae]]]]]].
As the families of Eupolypod I-clade could be seen as subfamilies of the large Polypoidaceae s.l., so the families of Eupolypod II-clade could be seen as subfamilies of the
widely distributed Aspleniaceae Newman nom.cons. Unlike many previous classifica-
Nat. Croat. Vol. 31(1), 2022
153
tions (also in our Garden), this “old” family in its “new” classification (according to
PPG-1, 2016) is monotypic: strictly speaking, it holds just the eponymous Asplenium (or
perhaps also, the not widely recognized Hymenasplenium and Hemidictyum) – needless
to say, a view which is highly disputed. This “supergenus” is indubitably paraphyletic,
including more than 40 formerly described genera, with ca 700 species (APweb, 2021
and literature within). Some of the characteristics common to these (very variable) “new
Aspleniums” are: fronds with decurrent margins and lateral ridges along rachis’; often
epiphytic and litter trapping plants.
To add to the data published by Heinz (1895-96) and Kovačić (2015), we have grown
different outdoor ‘spleenworts’ in the Garden for decades (Tab. 1). There are 20, often
lithophytic, Asplenium-taxa in Croatian flora, mostly under the older synonymy according to the botanic sources used in this part of Europe (extensive literature could be found
in FCD). For example, the original ‘Linnean’ name A. trichomanes-ramosum L. was at its
time widely accepted, but in “everyday use” was A. viride Huds., today valid for both
WFO and PoWO. Asplenium ceterach L. (PoWO, 2021; also FCD, 2021) is again a “more
popular” Ceterach officinarum Willd. (WFO, 2021), while A. scolopendrium L. was always
better known as Phyllitis scolopendrium (L.) Newman (in Heinz’s times Scolopendrium
phyllitis Roth). Statutorily strictly protected, the endemic A. hybridum (Milde) Bange is
nomen conservandum of Croatian flora (FCD, 2021; also PoWO, 2021), though Phylitopsis
hybrida (Milde) T.Reichst according to WFO (2021). In 2017, we grew some from spores
(Photo-tab. 21 in Suppl.) brought by Dr Vedran Šegota from the Island of Rab. Critically
endangered (CR), elusive A. sagittatum (DC.) Bange, which Dr Šegota also brought from
the single confirmed Croatian locality in the Island of Krk, sprouted very poorly and
lived just briefly.
Members of the “aspleniinian” family Cystopteridaceae Schmakov share long-creeping rhizomes; veins reach the frond margins; hood-like or unresisting indusium; n = 40.
To the recent views, three genera belong here (Heinz did not list any). PoWO does not
recognize this family but classifies its genera under Aspleniaceae. Cystopteris fragilis (L.)
Bernh. and Gymnocarpium robertianum (Hoffm.) Newman are Croatian native, which we
brought many times from the field research and planted in our rockeries with autochthonous species. Being the mountainous plants, they dislike hot and dry Zagreb summers, and withered soon. According to FCD, both genera belong to the Woodsiaceae
family, as does Athyrium (Athyriaceae), Matteucia (syn. Onoclea; Onocleaceae) and Woodsia (the single Croatian representative of the genus, W. ilvensis (L.) R.Br., was never inventoried in our collections). The modern view of the mostly montane, northern hemisphere ‘cliff-ferns’, Woodsiaceae Herter differs from the FCD’s: it (disputably) includes
a single genus (Woodsia, with up to 50 species; APweb, 2021), and it is a sister to the rest
of the ‘Aspleniinae’. PoWO does not recognize this monotypic family, but includes this
genus in Aspleniaceae. We grew three Woodsia-species in 2016 (Tab. 1), out of which
Woodsia plummerae Lemmon is the hardiest. Several examples of Cystopteridaceae and
Woodsiaceae can be seen in Photo-tab. 22 in Suppl.
The mostly terrestrial, understory Athyriaceae Alston encompasses 2(3-7) genera
with a few constant features by which they can be identified as members of this (disputed) family (for instance, sporangia have stalks two or three cells wide in the middle,
brown monolete spores). We are growing three Athyrium-taxa in our fernarium, out of
which the cosmopolite A. filix-femina (L.) Roth is a Croatian native (Tab. 1; Photo-tab.
23 in Suppl.). It is, traditionally, grown side-by-side with Dryopteris filix-mas (L.) Schott,
154 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
as the “pair” is in Latin, and many other languages, called the ‘lady fern’ (Croat. “ženska paprat”; Lat. filix-femina) and the ‘male fern’ (Croat. “muška paprat”; Lat. filix-mas).
This evokes the “Shakespearian” and much older times, when the world of ferns was
poorly understood (“We have the receipt of fern-seed, we walk invisible.” – “Henry IV, Part
1”, Act 2, Scene 1). Mature fronds of Athyriaceae are often abundant in anthocyanins
(APweb, 2021), a feature happily used in horticulture.
The “Aspleniinian” clade of [Blechnaceae + Onocleaceae]] is morphologically characterized by dimorphic (fertile and sterile) fronds, for which they are also popular in
horticulture. Northern-hemisphere Onocleaceae Pic.Serm. have four genera with five
species in total. Croatian native is Matteucia (Onoclea) struthiopteris (L.) Tod., grown in
our older collections, and flourishing in our fernarium for years (Photo-tab. 24 in Suppl.).
Onoclea struthiopteris was noted in the Registry of Spores Received in 1925, separately
from Matteucia struthiopteris Tod. and Struthiopteris germanica Willd., showing that the
synonymy was already complex to navigate. Onoclea sensibilis L. was inventoried in
1904 and again in 1963 (Tab. 1); we acquired some young plants in 2021. Fossils of this
species are known from Palaeocene North America (62-58 million of years ago), being
remarkably similar to the extant individuals.
Cosmopolitan Blechnaceae Newman have reddish young fronds (Photo-tab. 25 in
Suppl.) with linear sori. The family includes 24 genera (with many synonyms), out of
which we have (or had) in the Garden representatives of Blechnum, Doodia and Stenochlaena (Kovačić, 2015; Tab. 1). There is a significant discrepancy in affiliation species
and genera, according to PoWO and WFO. For example, our old S. tenuifolia (Desv.)
Moore (val. Lomariopsis tenuifolia (Desv.) Christ) according to PoWO) belongs to the
Polypodiaceae – not even Aspleniaceae – family. The acidophilic Blechnum spicant (L.)
Roth is the only member of this family in Croatian flora, grown on-and-off (with poor
success) for years. In 2016 we grew 3 subspecies’ of Blechnum penna-marina (Poir.) Kuhn,
Woodwardia orientalis Sw. and, after a long while, Blechnum brasiliense Desv. again, which
we had in our older collections (Tab. 1).
The terrestric Thelypteridaceae Pic.Serm. have an especially problematic range of
8 to 30 genera, with more than 1000 species (APweb, 2021). ‘Thelypteroid’ ferns were
often included in Dryopteris, and when you grow them side by side, you can see why:
morphologically, they could be very similar. However, there are many differences
between the two groups (as Dryopteris belong to “polypodiinean” and Thelypteris to
“aspleniinean” ferns), supported by genetic evidence, which shows that Thelypteridaceae are clearly monophyletic. However, the division of taxa into genera has been
described as “highly controversial and fluctuating” (Hassler & Schmitt, 2021); PoWO
does not recognize this family, and even places some genera into Polypodaceae. Consequently, it is really hard for me to say, at this moment, whether Heinz (1895-96) had
some representatives of ‘thelypterioids’ in the Garden collections, as the synonymy is
overwhelming. For example: we have had the Croatian native Thelypteris palustris
Schott (orig. Nephrodium thelypteris (L.) Strempel, Dryopteridaceae) in the Garden since
1962 (Photo-tab. 26 in Suppl.); the synonymy of this plant is utterly confusing (Tab. 1).
WFO does not recognize synonymy of the classic South-European taxa, proposed by
the first explorers of this part of Europe, whose works are fundamental to our local
and regional Floras (e.g.., Hayek, Visiani, Javorka, etc.). In 2016 we tried to grow several representatives of Thelypteridaceae for our fernarium (Tab. 1), with moderate success.
Nat. Croat. Vol. 31(1), 2022
155
CONCLUSIONS
According to the new and some old (recovered) sources, I have amended the historical data of the ferns growing in Botanical Garden since 1895 until the end of 2021
(Tab. 1). Though the synonymy of taxa is immense, and many of the old entries were
registered without the authors of plant names, my educated guess at this point is that
we had in our Garden collections around 376 species of ferns, including, presumably,
Heinz’s ca 35 in 1895, from 29 families. At this moment (December, 2021) we grow 86
taxa from 20 fern families, including Equisetaceae and Psilotaceae.
Annus Horribilis of 2020, which brought not only COVID-19 pandemic with lockdowns at planetary level, but also severe earthquakes and floods to Zagreb in particular, directly and indirectly diminished our collections. Consequently, among other
things, we lost a lot of ferns grown from spores and kept in our warm greenhouses,
due to the weather extremes and loss of heating while temperatures dropped below 0˚
C. The year 2021 was not much better.
Received January 10, 2022
ZAKLJUČCI
Prema novim i nekim starim (naknadno pronađenim) izvorima, dopunjeni su povijesni podaci o papratnjačama u zbirkama Botaničkog vrta između 1895. i kraja 2021.
(Tablica 1 u Dodatcima). Kako je sinonimika paprati nepregledna, a mnogi stari unosi
zabilježeni bez autora vrste, pretpostavljam da je kroz naše zbirke dosada prošlo oko
376 vrsta papratnjača, uključujući 35 Heinzovih iz 1895., iz oko 29 porodica. U ovom
trenutku (prosinac 2021.) uzgajamo 86 svojti iz 20 porodica, uključujući Equisetaceae
i Psilotaceae.
Annus Horribilis 2020. – koja nije donijela samo pandemiju bolesti COVID-19 s
prestankom rada na planetarnoj razini, nego i razorne potrese i velike poplave posebno Zagrebu – izravno i neizravno utjecala je i na naše zbirke. Posljedično, između ostalih vrsta, izgubili smo više svojta paprati uzgojenih iz spora i držanih u našim toplim
staklenicima, zbog vremenskih ekstrema i obustave grijanja u duljem razdoblju kad su
vanjske temperature pale ispod 0˚ C.
Godina 2021. nije bila puno bolja.
Primljeno 10. siječnja 2022
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Christenhusz, M.J.M. & Chase, M. W., 2018: PPG recognises too many fern genera. Taxon 67(3), 481–
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Engler, A. & Prantl, K. eds., 1887–: Die Natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten (in German; 33 parts in 23 volumes). Leipzig: W. Engelmann.
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GBIF Backbone Taxonomy. Gymnogramma gloriosa Hort. (in Tab. 1) https://www.gbif.org/es/species/6035185. Accessed via GBIF.org December 2021]
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8.20. [accessed December 2021]
Heinz, A., 1892: Über Scolopendrium hybridum Milde. Berichte der Deutschen Botanischen Gesellschaft 10,
413-422.
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Hidalgo, O., Pellicer, J., Christenhusz, M.J. M., Schneider, H. & Leitch, I.J., 2017: Genomic gigantism
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Moran, R.C. & Smith, A.R., 2018: Are there too many fern genera? Taxon 67(3), 473–480.
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Dinaric Mountains. Acta Botanica Croatica 54(1), 47-62.
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Photo-tables 6-26 are available in Supplement, only in online version of this article.
All photos were taken by the Author.
Nat. Croat. Vol. 31(1), 2022
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b
c
a
d
Photo-tab. 6: Early predecessors of horsetails: ab) Sphenophyllum alatifolium had large, whorled, and
sometimes deeply lobed leaves with dichotomous venation, suggesting that the tiny leaves of extant
Equisetum may be reduced megaphylls, rather than true microphylls. (Detail from the Garden exhibition in 2018). Famous c) Annularia stellata d) and Astrophyllites equisetiformis were already Equisetum-like
plants (Fossils from the Garden collection, originating from Spain).
158 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
a
b
c
d
Photo-tab. 7: Horsetails in the Garden: a) Equisetum arvense was subjected to various investigations
during the centuries; a1) non-photosynthetic strobili, produced early in spring. b) E. telmateia, with its
b1) “unkempt”, hair-like long leaves arranged in whorls. c) E. fluviatile is the first recorded horsetail in
our collection; c1) whorls of short ascending and spreading branches, c2) strobili on top of the photosynthetic branches. d) E. giganteum ‘El Tabacal’ with a backdrop of Matteucia struthiopteris in our
fernarium.
Nat. Croat. Vol. 31(1), 2022
159
b
a
c
Photo-tab. 8: a) Diorama of the Rhynie chert in Scotland, with home-made prototaxite and a geyser. b)
Detail of Devonic diorama with “rhinias”. c) Branch of Psilotum nudum, perfectly – though falsely –
imitating dichotomous branching of the earliest land plants.
160 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
a
b
c
d
Photo-tab. 9: Marattiales: a) Mature specimen of Angiopteris evecta in collection of ancient plants of the
Jardin botanique de la Ville de Paris. b) Gametophytes (prothalia) of A. evecta grown in our Garden
laboratory (sadly, never produced sporophytes). c) Polymorphopteris magdalenae and d) Seftenbergia grunerii, some of the extinct marattiaceous ferns in the Botanical Garden fossil collection.
a
b
Photo-tab. 10: Osmundales: a) Osmunda regalis lives happily in our Garden fernarium, as well as in
several other places outdoors, though never reaching the 3-meter-height, as in nature. b) In the Exhibition of 2018, the “royal fern” gets a special place (bottom left) in our little “Cretaceous garden”, with
other ancient plants from our collections.
Nat. Croat. Vol. 31(1), 2022
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b
c
d
a
Photo-tab. 11: Schizeales: a) Lygodium japonicum (Lygodiaceae) climbing from the hanging baskets of
the Garden’s Victoria house. Salviniales: b) Salvinia natans and Azolla filiculoides (Salviniaceae) together
in the Victoria house. c) Marsilea quadrifolia and d) Regnellidium diphyllum (Marsileaceae) are inhabitants
of the outdoor basins.
b
a
c
Photo-tab. 12: Cyatheaceae: a) Dicksonia antarctica in the Garden fernarium overwinters indoors. cd)
Cibotium regale, Dicksonia antarctica and Sphaeropteris (Cyathea) cooperi grown in our Garden laboratory.
162 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
b
a
c
Photo-tab. 13: Dennstaedtiaceae: a) Croatian native, Pteridium aquilinum lives in the Garden fernarium.
b) Hypolepis millefolium and c) Histiopteris incisa grown in our Garden laboratory, live indoors.
Nat. Croat. Vol. 31(1), 2022
a
163
b
c
e
f
d
i
h
g
k
l
j
Photo-tab. 14: Pteridaceae: The “maidenhair ferns” prefer moist, well-drained sites: a) Adiantum caudatum, b) A. macrophyllum, c) A. trapeziforme and d) Bolbitis heteroclita. So-called “cheilanthoid ferns”,
some 400 or more species, can grow in very dry conditions: e) Cheilanthes lanosa, f) Cryptogramma acrostichoides, g) Hemionitis palmata and h) Llavea cordifolia. Many are epiphytes, preferably grown in terreriums: i) Microgramma pilloseloides and j) Pellaea rotundifolia. Some species hyperaccumulate arsenic,
while other could be invasive: k) Pteris cretica 'Albolineata' and l) Pteris multifida.
164 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
a
b
c
d
f
e
Photo-tab. 15: Polypodiaceae 1: a-d) Data-cards of Microsorum punctatum and Niphidium (Campyloneurum) crassifolium, with many synonyms in WFO/PoWO databases each, changed their names in our
Garden database no fewer than six times. The synonymy between Polypodium-Phyamatodes-Selliguea-Microsorum-Cochlidium taxa is very hard to navigate – even worse is the synonymy of Polypodium-Phyamatodes-Pessopteris-Pleuririum-Niphidium-Campyloneurum. e) Microsorum punctatum and f) Niphidium (Campyloneurum) crassifolium.
Nat. Croat. Vol. 31(1), 2022
165
b
d
a
f
c
e
g
Photo-tab. 16: Polypodiaceae 2: Three taxa of bright green stag horns in our collection: a) Platycerium
bifurcatum, b) P. alcicorne and c) P. willinckii are epiphytic, as well as bluish d) Belvisia (Lepisorus) platyrhynchos and e) Phlebodium aureum ‘Umbellatum’. Two Croatian native polypodiums: f) P. vulgare and
g) P. cambricum might be similar in appearance, but differ in lifestyle (evergreen vs. deciduous).
166 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
b
c
a
d
Photo-tab. 17: Dryopteridaceae 1: Dryopteris: brightly coloured East-Asian a) D. erythrosora, Croatian
native b) D. filix-mas; American c) D. goldieana and subtropical d) D. pseudocaenopteris (Peranema aspidioides) with hairy young leaves, living in the greenhouse.
a
b
c
d
Photo-tab. 18: Dryopteridaceae 2.: Polystichum: Croatian native species (c) clearly differ during the
wintertime: a) P. setiferum is mostly deciduous, while b) P. aculeatum is evergreen. The famous ornamental d) P. setiferum ‘Plumosum Densum’ rarely overwinters with fronds.
Nat. Croat. Vol. 31(1), 2022
167
b
d
c
a
e
Photo-tab. 19: Davalliaceae: a) Davallia solida (‘Superba’), b) D. (Humata) pectinata, c) D. embolostegia
grown from spores. Tectariaceae: d) Tectaria (Quercifilix) zeylanica grown from spores, e) T. cicutaria
bearing button-like bulbils (propagules) at pinnae axils.
b
d
c
a
e
Photo-tab. 20: Nephrolepidaceae: a) Nephrolepis exaltata ‘Selecta’ lives in our collection since 1963, and
along the regular fronds it sprouts b), the feathery ones. c) Old N. exaltata planted permanently in our
fernarium. d) N. exaltata aff. ‘Curly Locks’ sprouted in 2016, among germinating spores. e) N. cordifolia
aff. 'Duffii', popular "Lemon Button Fern".
168 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
b
c
a
f
d
g
e
h
i
Photo-tab. 21: Aspleniaceae: abc) Asplenium ruta-muraria on the stone-walls of our Faculty building. d)
A. adiantum-nigrum and A. trichomanes-ramosum (A. viride) are sensitive species of Croatian flora, while
e) A. trichomanes is even frost-tolerant. f) A. scolopendrium lives in our fernarium “since forever”. g) A.
hybridum is a stenoendemic of NE Adriatic islands, grown from spores for our glasshouse collection,
where exotic spleenworts permanently live: for example, h) large A. nidus or i) invasive A. viviparum,
etc.
Nat. Croat. Vol. 31(1), 2022
169
c
a
b
d
Photo-tab. 22: Cystopteridaceae: a) Cystopteris tasmanica grown from spores. Woodsiaceae: b) young
plants grown from spores. c) Young Woodsia plummerae. d) Older individuals of W. plummerae planted
in our fernarium are deciduous.
a
b
c
Photo-tab. 23: Athyriaceae: a) Athyrium filix-femina growing side-by-side with its famous cultivar b)
'Frizelliae'. c) A. (Anisocampium) niponicum ‘Metallicum’, with beautifully coloured fronds.
170 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
a
c
e
b
d
f
Photo-tab. 24: Onocleaceae: Life cycle of Matteucia (Onoclea) struthiopteris: sterile fronds in ab) Spring,
and cd) Summer. ef) Young and mature fertile fronds.
a
b
d
c
e
f
Photo-tab. 25: Blechnaceae: a) Blechnum brasiliense grown from spores b) in 2016 have lovely reddish c)
young fronds. d) Croatian native, B. spicant is a characteristic species of dark fir forests (Blechno-Abietetum) in the mountain region. e) B. penna-marina grown from spores, as well as f) young Woodwardia
orientalis.
Nat. Croat. Vol. 31(1), 2022
171
b
a
c
d
Photo-tab. 26: Thelypteridaceae: a) Croatian native Thelypteris palustris. b) Th. cordata was purchased,
while from spores were grown c) Th. (Parathelypteris) beddomei and d) Th. kunthii.
Tab. 1.
WFO:
Order/clade & Family: EQUISETALES – Equisetaceae (PPG-1, 2016 via APweb, 2021)
Equisetum L.
Equisetum L.
Equisetum spp.
Equisetum arvense L.
Equisetum arvense L.
E. arvense L.
Equisetum fluviatile L.
Equisetum fluviatile L.
E. limosum
E. fluviatile L. (= E- limosum L."
Equisetum giganteum L. (cult.)
Equisetum giganteum L. (cult.) E. 'El Tabacal'
Equisetum hyemale L.
Equisetum hyemale L.
Equisetum hiemale L.
Equisetum pratense Ehrh.
Equisetum pratense Ehrh.
E. pratense Ehrh.
Equisetum sylvaticum L.
Equisetum sylvaticum L.
E. silvaticum L.
Equisetum telmateia Ehrh.
Equisetum telmateia Ehrh.
E. sylvaticum L.
E. telmateia Ehrh.
E. telmateia Ehrh.
Heinz,
1895-6
1904
– 1948
yes
1917
E. telmateia Ehrh.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
"various, by the Grotto"
Zagreb, Sava river banks
spreading locally
2018
1958
Zagreb Nursery 1961
Equisetum giganteum L. 'El
Vodnjan Nursery 2018
Tabacal'
Hand-written inventoy of
Medicinal plants
NP Medvednica 2021
Tabačka ravan (Bosnia &
Herc.) 1972
Zalesina 1977
Ivanščica Mt 2004
NP Žumberak 2018
Zagreb, Sava river banks
2021
Order/clade & Family: PSILOTALES – Psilotaceae (PPG-1, 2016 via APweb, 2021)
Psilotum nudum (L.) P. Beauv.
Psilotum nudum (L.) P. Beauv.
Psilotum nudum
ELTE Budapest
(Hungary) 2017
P. nudum (L.) P. Beauv.
Order/clade & Family: OPHIOGLOSSALES – Ophioglossaceae (PPG-1, 2016 via APweb, 2021)
Botrychium lunaria (L.) Sw.
Botrychium lunaria (L.) Sw.
B. lunaria Swartz
Ophioglossum vulgatum L.
Ophioglossum vulgatum L.
O. vulgatum L.
O. vulgatum L.
Order/clade & Family: MARATTIALES – Marattiaceae (PPG-1, 2016 via APweb, 2021)
Angiopteris evecta (G. Forst.)
Angiopteris evecta (G. Forst.)
Angiopteris longifolia
Hoffm.
Hoffm.
A. evecta Hoffm.
listed in Registry of spores,
1925/26
yes
NP Velebit 1975
NP Velebit 2013
yes
Vienna-Belvedere
(Austria) 2017
short living above-ground
short living above-ground
Several before, uninventorized
172 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Order/clade & Family: OSMUNDALES – Osmundaceae (PPG-1, 2016 via APweb, 2021)
Osmunda claytoniana L.
Osmunda claytoniana L.
O. claytoniana
Osmunda regalis L.
Osmunda regalis L.
Osmunda regalis
O. regalis
O. regalis L.
sprouted in 1926 (register of
spores)
1926
1904
"outdoors"
yes
Borlin, Karlovac 1962
Glina, Banovina 2016
O. regalis L.
Osmunda spectabilis Willd.
Osmunda gracilis Link (ambig.) O. gracile
Osmundastrum cinnamomeum Osmundastrum
Osmunda cinnamomea
(L.) C.Presl
cinnamomeum (L.) C.Presl
Todea Willd.
Todea Willd. ex Bernh.
Todea spp.
Order/clade & Family: HYMENOPHYLLALES – Hymenophyllaceae (PPG-1, 2016 via APweb, 2021)
Hymenophyllum
Hymenophyllum
Hymenophylum tunbridgense
tunbrigense (L.) Sm.
tunbrigense (L.) Sm.
Vandenboschia radicans (Sw.)
Trichomanes radicans Sw.
Trichomanes radicans
Copel.
Order/clade & Family: SCHIZEALES – Anemiaceae (PPG-1, 2016 via APweb, 2021)
Anemia phyllitidis (L.) Sw.
Anemia phyllitidis (L.) Sw.
Aneimia Phyllitidis
(Schizeaceae)
Anemia mexicana Klotzsch
Anemia mexicana Klotzsch
A. mexicana Klotzsch
(Schizeaceae)
Anemia rotundifolia Schrad.
Anemia rotundifolia Schrad.
A. rotundifolia Schrad.
(Schizeaceae)
Order/clade & Family: SCHIZEALES – Lygodiaceae (PPG-1, 2016 via APweb, 2021)
Lygodium japonicum (Thunb.) Lygodium japonicum (Thunb.)
Lygodium japonicum (Sw.)
Sw. (Schizeaceae)
Sw.
L. japonicum (Thunb.) Sw.
1904
yes
extinct in Croatia
yes
yes
yes
1904
191?
München (Germany)
2016
yes
1904
191?
yes
Vienna-Belvedere
(Austria) 2016
Ligodium palmatum
1922
1932
173
Lygodium palmatum (Bernh.)
Sw.
in 2018: new plants from those
spores
1904
L. japonicum (Thunb.) Sw.
Lygodium palmatum (Bernh.)
Sw. (Schizeaceae)
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Order/clade & Family: SALVINIALES – Salviniaceae (PPG-1, 2016 via APweb, 2021)
Azolla Lam.
Azolla Lam.
Asolla
Azolla caroliniana Willd.
Azolla caroliniana Willd.
Azolla caroliniana Willd.
Azolla caroliniana Willd.
A. filiculoides Lam.
A. filiculoides Lam.
Salvinia auriculata Aubl.
Salvinia auriculata Aubl.
Heinz,
1895-6
1904
– 1948
yes
1920
1948
Salvinia minima Baker
Salvinia natans (L.) All.
Salvinia natans (L.) All.
yes
1948
Marsilea hirsuta R. Br.
Marsilea macropoda Engelm. ex Marsilea macropoda Engelm.
A. Braun
ex A. Braun
Marsilea quadrifolia L.
Marsilea quadrifolia L.
Marsilea strigosa Willd.
M. hirsuta R. Br.
Marsilia macropoda Engelm.
Marsilia quadrifolia
M. quadrifolia L.
Marsilea pubescens Ten.
Marsilia pubescens Ten.
(ambig.)
Marsilea strigosa Willd. (amb.) M. strigosa Willd.
*changed to S. molesta
D.S.Mitch. acc. to TROPICOS
Living specimen
"fam. Marsiliaceae"
1961
Strassbourg (France)
1965
Göttingen (Germany)
1969
Göttingen (Germany)
1969
Strassbourg (France)
1965
Marsilea drummondii A. Br.
Marsilea hirsuta R. Br.
Berlin-Dahlem
(Germany) 2013
"from Osijek vicinity"
Oslo 1948
1961
yes
Marsilia drummondii A. Br.
Hand-written inventory
Living specimen
Hand-written inventory
1963
S. minima Baker
Salvinia natans
S. natans All.
S. natans Hoffm.
Order/clade & Family: SALVINIALES – Marsileaceae (PPG-1, 2016 via APweb, 2021)
Marsilea L.
Marsilea L.
Marsilia
Marsilea drummondii A. Braun Marsilea drummondii A. Braun Marsilia drummondii A. Br.
Oslo 1948
1961
Basel (Germany) 1971
1920
1938
S. auriculata
S. auriculata Aubl.*
Salvinia minima Baker
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
yes
1961
1961
Padova (Italy)1985
reconstructed card
174 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
Regnellidium diphyllum
Regnellidium diphyllum
Regnellidium diphyllum
Lindm.
Lindm.
Lindm.
Order/clade & Family: CYATHEALES – Cibotiaceae (PPG-1, 2016 via APweb, 2021)
unknown sp.* (gen. Cibotium = Cibotium regale Verschaff. &
Cibotium Regale
fam. Cyatheaceae)
Lem.
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Bonn (Germany) 2021
191?
Tübingen (Germany)
2016
C. regale Verschaff. & Lem.
Cibotium schiedei Schltdl. &
Cibotium schiedei Schltdl. &
Cibotium Schiedei
Cham.
Cham.
Order/clade & Family: CYATHEALES – Cyatheaceae (PPG-1, 2016 via APweb, 2021)
Alsophila R.Br.
Alsophila R.Br.
Alsophila
Alsophila australis R.Br.
Alsophila australis R.Br.
Alsophylla australis Br.
(ambig.)
*probably from horticulture
(syn. "Cibotium spectabile
(Hort.)")
191?
yes
1904
191?
1925
Alsophila australis
A. australis R.Br.
(in Kovačić, 2015 as Cyathea
australis Domin)
p.p.
yes
Alsophila leichhardtiana F.
Muell.
Cyathea dealbata (G.Forst.) Sw.
yes
191?
yes
191?
Brno (Czech Republic)
2016
Tübingen (Germany)
2016
191?
yes
marked with "?"
"fam. Cyatheaceae"
175
Cyathea australis Domin (amb.) C. australis Domin
Alsophila leichhardtiana F.
A. Leichhardtiana F.Muell.
Muell (ambig.)
Cyathea dealbata Sw. (ambig.) C. dealbata
C. dealbata Sw.
Gymnosphaera glabra Blume
Cyathea glabra (Blume) Copel. Alsophila glabra
Sphaeropteris cooperi (F. Muell.) Sphaeropteris cooperi (F.
S. cooperi (F. Muell.) R.M. Tryon
R.M. Tryon
Muell.) R.M. Tryon
Cyathea cooperi (Hook. ex F.
Muell.) Domin
Sphaeropteris myosuroides
Alsophila aurea Fée (ambig.)
Alsophila aurea
(Liebm.) R.M.Tryon
Order/clade & Family: CYATHEALES – Dicksoniaceae (PPG-1, 2016 via APweb, 2021)
Dicksonia L'Hér. (Cyatheaceae) Dicksonia L'Hér.
Dicksonia
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
PoWO (Kew):
WFO:
Dicksonia antarctica Labill.
Balantium antarcticum (Labill.)
Dicksonia antarctica (Labill.)
C. Presl
Heinz,
1895-6
1904
– 1948
191?
1932
D. antarctica Labill.
D. antarctica Labill.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
also sprouted in 1926 (register
yes
of spores)
Tübingen (Germany)
2016
bought, with a permit of
Zagreb Nursery 2021
Australian government
Order/clade & Family: POLYPODIALES – Aspleniaceae (PPG-1, 2016 via APweb, 2021)
Asplenium L.
Asplenium L.
Asplenium spp.
yes
Asplenium L.
Ceterach Willd.
Ceterach spp.
yes
Asplenium L. (p.p.)
Asplenium L. (p.p.)
Scolopendrium spp.
yes
191?
Asplenium adiantum-nigrum L. Asplenium adiantum-nigrum L. A. adiantum-nigrum L.
A. adiantum-nigrum L.
A. adiantum-nigrum L.
A. adiantum-nigrum L.
Naples (Italy) 1959
Vršac (Serbia) 1965
Lim channel 1966
Koločep 1970
Matka (North
Macedonia) 1973
Orjen Mt (Montenegro)
1978
NP Velebit 2003
NP Učka 2016
brought as a living plant
in hand-written inventory of
1955
glasshouse in 1955
"Palaeotropic Asplenium
species' with bulbils atop of
their fronds", fam.
Polypodiaceae
A. adiantum-nigrum L.
A. adiantum-nigrum L.
Asplenium australasicum Hook. Asplenium nidus L.
Asplenium bulbiferum G. Forst.
Asplenium bulbiferum G.
Forst.
A. adiantum-nigrum L.
A. adiantum-nigrum L.
A. nidus var. australasicum
Hook.
(?)
A. "bulbi/forme"
"By the Grotto", fam.
Polypodiaceae
"By the Grotto", fam.
Polypodiaceae
"By the Grotto", fam.
Polypodiaceae, probably
Croatian native species
yes
1904
191?
1932
(perhaps of some other
bulbiform taxon)
176 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Asplenium ceterach L.
Ceterach officinarum Willd.
C. officinalis
yes
C. officinalis Willd. var.
crenatum Moore
yes
Heinz,
1895-6
1904
– 1948
A. bulbiferum Forst.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
C. officinarum DC.
Asplenium ceterach L.
A. ceterach L.
A. ceterach L.
A. ceterach L.
A. ceterach L.
A. ceterach L.
177
A. ceterach L.
A. ceterach L.
A. ceterach L.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
yes
in hand-written inventories in
1950-ies
Orig. author (Willd.) was
"Alps" 1950
changed to"DC." acc. to
"Analytic Flora of Yugoslavia"
Mali Lošinj 1955
Osor 1962
Mosor Mt 1962
Svilaja Mt 1963
Dinara Mt 1963
NP Plitvice 1964
Vis 1964
NP Medvednica 1965
Kopaonik (Serbia) 1965
Lišani 1967
NP Mljet 1968
Orebić 1969
Orjen Mt (Montenegro)
1970
Pelješac 1971
Sućeška (Bosnia & Herc.)
1973
Đerdap (Serbia) 1975
NP Biokovo 1975
Mosor Mt 1977
Lovćen Mt (Montenegro)
1978
Brač 1978
Koločep 1979
NP Mljet 1979
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
A. ceterach L.
A. ceterach L.
A. ceterach L.
A. ceterach L.
Asplenium daucifolium Lam.
Asplenium viviparum (L. f.) C.
Ceanopteris (?) viviparum
Presl
Asplenium dimorphum Kunze
Asplenium dimorphum Kunze
A. dimorphum Kunze
(ambig.)
1904
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Olib 1983
NP Učka 2005
Murter 2010
Konavle 2011
probably (illegible)
Caenopteris vivipara
P.J.Bergius
yes
Berlin-Späth (Germany)
Living specimen
2021
NP Risnjak 1965
NP Velebit 1975
NP Velebit 2003
A. dimorphum Kunze
Asplenium fissum Kit.
Asplenium hybridum (Milde)
Bange
Asplenium fissum Kit. (ambig.) A. fissum Kit. ex Willd.
A. fissum Kit. ex Willd.
A. fissum Kit. ex Willd.
Phyllitopsis hybrida (Milde) T.
Scolopendrium hybridum
Reichst.
Asplenium hybridum (Milde)
Bange
A. hybridum (Milde) Bange
Phyllitis hybrida (ambig.)
Phyllitis hybrida
Asplenium marinum L.
Asplenium sulcatum Lam.
Asplenium marinum L.
Asplenium nidus L. (cult.?)
Asplenium nidus L. (cult.?)
A. "Nidus A/avis"
Asplenium nidus L. (cult.)
Asplenium nidus L. (cult.)
A. nidus L.
A. nidus L. ‘Osaka’
Asplenium ruta-muraria L.
Asplenium ruta-muraria L.
A. ruta muraria
A. ruta-muraria L.
A. ruta-muraria L.
“from Lošinj island”
yes
fam. Polypodiaceae
Olib 2015
Rab 2016
191?
1932
1904
1904
191?
1932
(grown from spores)
Croatian subendemic
Probably this species; acc. To
Parey, A. Nidus-avis is a
culton (hort.)
yes
yes
in hand-written inventories of
glasshouse in 1950-ies
yes
NP Učka 1963
NP Velebit 1964
178 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
A. ruta-muraria L.
Asplenium sagittatum (DC.)
Bange
Asplenium scolopendrium L.
Asplenium scolopendrium L.
subsp. scolopendrium
Asplenium scolopendrium L.
(cult.)
Asplenium sagittatum (DC.)
Bunge
Asplenium scolopendrium L.
Scolopendrium officinale DC.
(ambig.)
Asplenium scolopendrium L.
(cult.)
A. sagittatum (DC.) Bange
Scolopendrium spp.
Phyllitis scolopendrium (L.)
Newman
Ph. scolopendrium (L.) Newman
Ph. scolopendrium (L.) Newman
Asplenium scolopendrium L.
A. scolopendrium L.
Scolopendrium officinale
1958
Jasenak 1971
Strahinjčica Mt 1971
Kašina 1993
NP Žumberak 2001
1904
A. septentrionale (L.) Hoffm.
yes
probably A. scolopendrium
'Undulatum' (group)
1904
Vršac (Serbia) 1965
reconstructed card
179
Asplenium septentrionale (L.)
Hoffm.
Rab 2017
yes
A. scolopendrium L. 'Crispum
Cristatum’
A. undulatum
Asplenium septentrionale (L.)
Hoffm.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Jajce (Bosnia & Herc.)
1966
Pelješac 1969
Martin Brod 1969
Pelješac 1971
Soteska (Slovenia) 1971
NP Žumberak 1972
Prokletije (Montenegro)
1973
Lovćen (Montenegro)
1978
Ravna Gora 1988
NP Velebit 2003
unknown
spreads locally
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Asplenium stellatum Colla
Asplenium fernandezianum
Kunze (ambig.)
Asplenium fernandezianum
Asplenium trichomanes L.
Asplenium trichomanes L.
A. trichomanes
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
Asplenium viride Huds.
Asplenium viride Huds.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. trichomanes L.
A. viride Huds.
A. viride Huds.
A. viride Huds.
A. viride Huds.
A. trichomanes-ramosum L.
A. trichomanes-ramosum L.
A. trichomanes-ramosum L.
A. trichomanes-ramosum L.
A. trichomanes-ramosum L.
Heinz,
1895-6
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
in hand-written inventories of
glasshouse in 1950-ies
yes
Klek 1961
Bohinj (Slovenia) 197
NP Plitvice 1964
Badanj 1964
Vršac (Serbia) 1965
Lim channel 1966
Pelješac 1969
Matka (North
Macedonia) 1973
Bled (Slovenia) 1971
Strahinjčica Mt 1979
NP Mljet 1982
Plomin 1982
Olib 1983
Samobor 2002
Vukova Gorica 2008
NP Velebit 1959
Bijele stijene 1961
Troglav (Slovenia) 1963
Badanj 1964
Bled (Slovenia) 1971
Samarske stijene 1972
Strahinjčica Mt 1973
NP Velebit 1973
Prokletije Mt
(Montenegro) 1973
180 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
A. trichomanes-ramosum L.
A. trichomanes-ramosum L.
A. trichomanes-ramosum L.
A. trichomanes-ramosum L.
unknown (A. viviparum Blume Asplenium viviparum (L.f.) C.
(?)
– val. A. daucifolium Lam.)
Presl
yes
Caenopteris vivipara
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Julian Alps (Slovenia)
1982
Savica 1985
Kostel 1990
Dubovac, Karlovac 2016
"Palaeotropic Asplenium
species' with bulbils atop of
yes
their fronds" fam.
Polypodiaceae
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
1904
Asplenium lineatum var.
viviparum (syn. Darea vivipara)
unknown combination; in
hand-written inventores of
glasshouse in 1950-ies
yes
Order/clade & Family: POLYPODIALES – Athyriaceae (PPG-1, 2016 via APweb, 2021)
Athyrium Roth (Aspleniaceae)
Athyrium Roth
Athyrium
Athyrium "Pri(t/r?)churense
cristata"
Athyrium filix-femina (L.) Roth Athyrium filix-femina (L.) Roth A. filix-femina (L.) Rth.
A. filix-femina (L.) Roth
A. filix-femina (L.) Roth
Athyrium filix-femina (L.) Roth Athyrium filix-femina (L.) Roth
Athyrium crispatum
(cult.)
(cult.)
Athyrium Frizeliae
yes
1904
Samarske stijene 1963
Strahinjčica Mt 1973
Sunger 2010
"By the Grotto" fam.
Polypodiaceae
unknown (illegible
handwriting)
FCD: Woodsiaceae
probably Athyrium
filix-femina 'Crispata'
1904
1904
Buzet Nursery 2006
val. Athyrium filix-femina (L.)
Roth 'Frizelliae'
Zagreb Nursery 2021
Athyrium niponicum (Mett.)
Hance (cult.)
Anisocampium niponicum
(Mett.) Y.C.Liu, W.L. Chiou &
M. Kato (cult.)
Athyrium niponicum
'Metallicum'
Buzet Nursery 2006
(?)
(?)
Athyrium goringianum pictum
1904
val. Athyrium filix-femina (L.)
Roth 'Frizelliae'
val. Athyrium niponicum
(Mett.) Hance 'Metallicum'
Perpaps A. niponicum var.
Pictum 'Metallicum'
181
A. 'Fritzeliae'
Tab. 1. Continued
WFO:
Heinz,
1895-6
Diplazium celtidifolium Kunze
Diplazium celtidifolium Kunze Asplenium celtidifolium
(Aspleniaceae)
Order/clade & Family: POLYPODIALES – Blechnaceae (PPG-1, 2016 via APweb, 2021)
Blechnum L. (Aspleniaceae)
Blechnum L.
Blechnum
Blechnum brasiliense Desv.
Blechnum brasiliense Desv.
B. brasiliense
1904
– 1948
1904
"By the Grotto", fam.
Polypodiaceae
yes
1904
191?
B. brasiliense Desv.
B. brasiliense Desv.
Blechnum castaneum (Makino)
Makino & Nemoto
Blechnum gibbum (Labill.) Mett.
Blechnum gibbum (Labill.) Mett.
? (cult.)
Blechnum castaneum Makino
& Nemoto (ambig.)
Lomaria gibba Labill. (ambig.)
Blechnum gibbum Mett.
(ambig.; ? cult.)
Blechnum hastatum Kaulf.
Blechnum hastatum Kaulf.
Blechnum microphyllum
(Goldm.) C.V. Morton
Blechnum moorei C.Chr.
Blechnum neohollandicum
Christenh.
Blechnum occidentale L.
Blechnum penna-marina
(Maxon & C.V.Morton) Kuhn
subsp. alpinum (R.Br.)
T.C.Chambers & P.A.Farrant
Blechnum penna-marina
(Maxon & C.V.Morton) Kuhn
subsp. microphyllum (Goldm.)
T.C.Chambers & P.A.Farrant
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
yes
B. castaneum Makino
Lomaria Gibba
fam. Polypodiaceae
yes
Blechnum 'Silver Lady'
Blechnum "pastatum v.
minimum" (? Perhaps B.
hastatum?)
Blechnum penna-marina (Poir.) B. penna-marina subsp.
Kuhn
microphyllum
Blechnum moorei C.Chr.
B. moorei C.Chr.
(ambig.)
Doodia aspera R.Br. (ambig.)
Doodia aspera R.Br.
Blechnum occidentale L.
Blechnum occidentale
1955
Riga (Latvia) 2016
Chemnitz (Germany)
2016
B. gibbum 'Silver Lady'
yes
probably was misspelle:
crossed and weitten over as "B.
occidentalis"
191?
Chemnitz (Germany)
2016
yes
yes
1954
191?
Blechnum penna-marina (Poir.)
B. penna-marina subsp. alpina
Kuhn (unknown subsp.)
Chemnitz (Germany)
2016
Blechnum penna-marina (Poir.) B. penna-marina subsp.
Kuhn (unknown subsp.)
microphylla
Chemnitz (Germany)
2016
as "Doodya" in older records
182 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
Heinz,
1895-6
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
PoWO (Kew):
WFO:
Blechnum penna-marina
(Maxon & C.V.Morton) Kuhn
subsp. penna-marina
Blechnum penna-marina (Poir.) B. penna-marina subsp.
Kuhn (unknown subsp.)
penna-marina
Chemnitz (Germany)
2016
Blechnum spicant (L.) Roth
Blechnum spicant (L.) Sm.
Bijele stijene 1961
B. spicant (L.) Roth*
Zalesina 1963
Štirovac 1964
Strahinjčica Mt 1973
Kupa 2000
Delnice 2004
Sunger 2010
Samobor 2018
Sunger 2018
Zagreb Nursery 2021
1904
1904
yes
yes
Vienna-Belvedere
(Austria) 2016
in hand-written inventories of
Karstic rockery in 1950-ies
yes
yes
yes
"Alps" 1949
Medak 1959
Bijele stijene 1961
reconstructed card; acc. to
FCD belongs to Woodsiaceae
183
B. spicant (L.) Roth
B. spicant (L.) Roth
B. spicant (L.) Roth
B. spicant (L.) Roth
B. spicant (L.) Roth
B. spicant (L.) Roth
B. spicant (L.) Roth
B. spicant (L.) Roth
B. spicant (L.) Roth
Blechnum spinulosum Poir.
Doodia caudata (Cav.) R. Br.
Do(o)dia caudata Br.
Woodwardia caudata
Lomariopsis tenuifolia (Desv.) Stenochlaena tenuifolia (Desv.) Stenochlaena tenuifolia (Desv.)
Christ (Polypodiaceae)
Moore
Moore
Stenochlaena palustris (Burm. f.) Stenochlaena palustris (Burm.
S. palustris Bedd.
Bedd. (Aspleniaceae)
f.) Bedd.
Woodwardia orientalis (Sw.) Sw.
Woodwardia orientalis Sw.
W. orientalis Sw.
(Aspleniaceae)
Order/clade & Family: POLYPODIALES – Cystopteridaceae (PPG-1, 2016 via APweb, 2021)
Cystopteris alpina (Lam.) Desv.
Cystopteris alpina (Jacq.) Desv. Cystopteris alpina
(Aspleniaceae)
Cystopteris bulbifera (L.) Bernh. Cystopteris bulbifera (L.)
C. bulbifera (L.) Bernh.
(Aspleniaceae)
Bernh.
Cystopteris fragilis (L.)
Cystopteris fragilis (L.) Bernh. C. fragilis (L.) Bernh.
Bernh. (Aspleniaceae)
C. fragilis (L.) Bernh.
C. fragilis (L.) Bernh.
*(never lived longer than two
years)
Nat. Croat. Vol. 31(1), 2022
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
C. fragilis (L.) Bernh.
C. fragilis (L.) Bernh.
C. fragilis (L.) Bernh.
C. fragilis (L.) Bernh.
C. fragilis (L.) Bernh.
C. fragilis (L.) Bernh.
C. fragilis (L.) Bernh.
C. fragilis (L.) Bernh.
Cystopteris tasmanica Hook.
(Aspleniaceae)
Gymnocarpium robertianum
(Hoffm.) Newman
(Aspleniaceae)
Cystopteris tasmanica Hook.
(ambig.)
C. tasmanica Hook.
Nephrodium robertianum
Prantl (ambig.)
Nephrodium robertianum
(Hoffm.) Prantl
N. robertianum (Hoffm.) Prantl
Gymnocarpium robertianum
(Hoffm.) Newman
G. robertianum (Hoffm.)
Newman
Order/clade & Family: POLYPODIALES – Davalliaceae (PPG-1, 2016 via APweb, 2021)
Davallia Sm. (Polypodiaceae)
Davallia Sm.
Davallia
Davallia denticulata (Burm. f.) Davallia denticulata (Burm. f.)
D. denticulata (Burm.) Mett.
Mett.
Mett. ex Kuhn
Davallia embolostegia Copel.
Davallia embolostegia Copel.
D. embolostegia Copel.
(ambig.)
Humata heterophylla (Sm.)
Davallia heterophylla Sm.
Humata heterophylla
Desv.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
NP Risnjak 1965
Troglav Mt (Slovenia)
1963
NP Učka 1973
Strahinjčica Mt 1973
Vrbas (Bosnia & Herc.)
1966
Idrijca (Slovenija) 1967
Donačka gora (Slovenia)
1988
NP Velebit 2003
Chemnitz (Germany)
2016
NP Učka 1963
NP Risnjak 1965
Gymnocarpium robertianum
(Hoffm.) Newman
Davallia pectinata Sm.
Davallia pectinata Sm.
Humata pectinata (Sm.) Desv.
FCD: Woodsiaceae
Bonn (Germany) 1994
NP Velebit 2003
fam. Polypodiaceae
yes
yes
Vienna-Belvedere
(Austria) 2016
191?
München (Germany)
2016
184 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Davallia solida (G. Forst.) Sw.
Davallia solida (G. Forst.) Sw.
D. solida Sw.
Davallia solida (G. Forst.) Sw.
Davallia solida Sw. ‘Superba’
(cult.)
Humata griffithiana (Hook.) C.
Davallia tyermannii (T.Moore)
Chr. var. tyermannii (T. Moore) H. tyermannii Moore
H.J.Veitch
Tagawa (ambig.)
H. tyermannii Moore
H. tyermannii Moore
Order/clade & Family: POLYPODIALES – Dennstaedtiaceae (PPG-1, 2016 via APweb, 2021)
Dennstaedtia adiantoides
Dennstaedtia adiantoides
Denstaedtia adiantoides
(Willd.) T.Moore
(Willd.) T.Moore
two different authors to
Dennstaedtia cicutaria (Sw.) T.
Dicksonia cicutaria
Dicksonia cicutaria
Moore
Heinz,
1895-6
1904
– 1948
Davallia solida (G. Forst.) Sw.
(cult.)
Dennstaedtia punctilobula
(Michx.) T.Moore
Histiopteris incisa (Thunb.) J.
Sm.
Hypolepis millefolium Hook.
Dennstaedtia punctilobula
(Michx.) T. Moore (unknown
syn.)
Histiopteris incisa (Thunb.) J.
Sm.
Hypolepis millefolium Hook.
(ambig.)
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Tübingen (Germany)
yes
1971
yes
Antwerp (Belgium) 1973 *incorrect (Tectaria cicutaria)
Antwerp (Belgium) 1980
Zagreb Nursery 2019
191?
191?
"Denstadenia cicularis"/
"Denstadmia sicularis"?
1932
illegible handwriting: perhaps
misspelled "Dennstaedtia
cicutaria"
Dicksonia punctilobula
1926
sprouted in 1926 (register of
spores)
Vienna-Belvedere
(Austria) 2016
Chemnitz (Germany)
2016
H. incisa (Thunb.) J. Sm.
H. millefolium Hook.
Microlepia cult.
Microlepia cult.
Microlepia hirta cristata
(probably cultivar, acc. to older
hort. Literature)
Microlepia speluncae (L.) T.
Moore
Microlepia speluncae (L.) T.
Moore
Davallia speluncae
1904
1904
M. hirta and M. cristata are
separate taxa: ambig. (WFO)
or with separate valid names
(PoWO)
yes
Cesargrad Mt 1969
FCD: Hypolepidaceae
185
M. speluncae (L.) Moore
Pteridium aquilinum (L.) Kuhn Pteridium aquilinum (L.) Kuhn P. aquilinum (L.) Kuhn
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
P. aquilinum (L.) Kuhn
P. aquilinum (L.) Kuhn
Order/clade & Family: POLYPODIALES – Didymochlaenaceae (PPG-1, 2016 via APweb, 2021)
Didymochlaena Desv.
Didymochlaena
Didimochlaena
(Polypodiaceae)
(Hypodematiaceae)
Didymochlaena truncatula (Sw.) Didymochlaena truncatula
Didymochlaena sinuata Desv.
J.Sm.
(Sw.) J.Sm.
Order/clade & Family: POLYPODIALES – Dryopteridaceae (PPG-1, 2016 via APweb, 2021)
Arachniodes aristata (G. Forst.)
Tindale (Polypodiaceae)
Arachniodes aristata (G. Forst.) Polystichum aristatum Swartz.
Tindale
(unknown author*)
Bolbitis heteroclita (C. Presl)
Ching (Polypodiaceae)
Cyrtomium falcatum (L. f.) C.
Presl (Polypodiaceae)
Bolbitis heteroclita (C. Presl)
Ching
Cyrtomium falcatum (L. f.) C.
Presl
Heinz,
1895-6
fam. Polypodiaceae
as syn. of D.sinuosa Desv.
Mendoza et al., 1999
1904
yes
C. falcatum L. fil.
yes
C. falcatum L. f.
‘Rochfordianum’
Cyrtomium fortunei J.Sm.
Cyrtomium Fortunei
Cyrtomium lonchitoides
(Christ) Christ
Polystichum lonchitoides
(Christ) Diels
Dryopteris Adans. (p.p.)
Nephrodium
*from basionym Aspidium
aristatum (G. Forster) Sw.
(Swartz), fam. Aspidiaceae?
yes
B. heteroclita (C. Presl) Ching
(Lomariopsidaceae)
Cyrtomium falcatum (L. f.) C.
Presl (cult.)
Dryopteris carthusiana (Vill.)
H.P. Fuchs
Dryopteris carthusiana (Vill.)
H.P. Fuchs
Dryopteris dilatata (Hoffm.)
A.Gray
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
NP Medvednica 1985
Rudanovac 2015
yes
Polystichum falcatum (L.f.) Diels
Cyrtomium falcatum (L. f.) C.
Presl (cult.)
Cyrtomium fortunei J.Sm.
(Polypodiaceae)
Cyrtomium lonchitoides (Christ)
Christ (Polypodiaceae)
Dryopteris Adans. (p.p.;
Polypodiaceae)
Dryopteris carthusiana (Vill.)
H.P. Fuchs
Dryopteris carthusiana (Vill.)
H.P. Fuchs var. carthusiana
Dryopteris dilatata (Hoffm.)
A.Gray
1904
– 1948
val. C. falcatum (L. f.) C.Presl
also in hand-written
inventories of glasshouse in
1950-ies
1932
yes
1904
yes
in hand-written inventories of
glasshouse in 1950-ies
yes
fam. Polypodiaceae
yes
D. carthusiana (Vill.) Fuchs
Strahinjčica Mt 1973
Aspidium "spin.(ulosum?)
interm.(edium?)"
1904
Aspidium dilatatum
1904
1975
Probably this species
186 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
PoWO (Kew):
WFO:
Dryopteris erythrosora (D.C.
Eaton) Kuntze
Dryopteris filix-mas (L.) Schott
Dryopteris erythrosora (D.C.
Dryopteris Erythrosora
Eaton) Kuntze
Dryopteris filix-mas (L.) Schott Aspidium filix-mas
Heinz,
1895-6
1904
– 1948
yes
1904
Nephrodium Filix mas
1917
"Dipteris filis mas" (?)
1932
Nephrodium filix-mas (L.) Rich.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
val. D. erythrosora (D.C.
Zagreb Nursery 2018
Eaton) Kuntze
Strahinjčica Mt 1973
yes
Dryopteris filix-mas (L.) Schott
Dryopteris filix-mas (L.) Schott
(cult.)
Dryopteris goeringiana (Kunze)
Koidz.
Dryopteris filix-mas (L.) Schott
Dryopteris cult.
(cult.)
Athyrium goeringianum
Athyrium goringianum
(Kunze) T. Moore
Zagreb Nursery 2021
(small cult.)
1904
Athyrium goringianum pictum
Perhaps
Athyrium niponicum 'Pictum'
1904
Tübingen (Germany)
2016
original card is lost; remaind a
small card with this
synonymy
yes
yes
1904
Mosor Mt 1977
yes
fam. Polypodiaceae
187
Dryopteris goldieana (Hook. ex Dryopteris goldiana (Hook. ex D. goldiana (Hook. ex Goldie) A.
Goldie) A.Gray
Goldie) A. Gray
Gray
"Nephrodium villarii (Bell) Beck
Dryopteris pallida (Bory) Maire
subsp. pallida (Bory) Hayek =
unknown
& Petitm.
Dryopteris pallida (Bory) C.
Chr. ex Maire et Petitmengin"
Dryopteris pseudocaenopteris Peranema aspidioides (Blume)
Diacalpe aspidioides Blume
(Kunze) Li Bing Zhang
Mett.
Dryopteris pseudosieboldii
Aspidium sieboldii Van Houtte
Aspidium Sieboldi van Houtte
Hayata
ex Mett. (ambig.)
Dryopteris villarii (Bellardi)
Dryopteris midshelkensis
D. villarii (Bellardi) Woynar
Woyn. ex Schinz & Thell.
Pavlov
Elaphoglossum crinitum (L.)
Chrysodium crinitum Mett.;
Chrysodium crinitum
Christ (Polypodiaceae)
Fil. (ambig.; Pteridaceae)
Hand-written inventoy of
Medicinal plants
unknown combination: I
presume it was this taxon
(Dipteris (Gleicheniales) is
native to tropical Asia)
noted as growing in Karstic
rockery in the 1950-ies
from other localities,
unregistered, spreads locally
Nat. Croat. Vol. 31(1), 2022
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Polybotrya sp. (Polypodiaceae)
Polybotrya sp.
Polybotrya numina/minima (?)
1904
Polybotrya minima (?)
191?
1932
Polystichum sp. (Polypodiaceae) Polystichum sp.
Polystichum "grandidentatum"
1904
Polystichum aculeatum (L.)
Roth
P. lobatum (Huds.) C. Presl var.
longilobum Milde
P. lobatum var. auriculatum
P. lobatum (Huds.) Presl.
Polystichum aculeatum (L.)
Roth ex Mert.
Heinz,
1895-6
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
illegible handwriting, both
unknown
unknown species
?
1958
1962
Troglav Mt (Slovenia)
1963
yes
P. lobatum (Huds.) Presl.
Polystichum braunii (Spenn.)
Fée
Polystichum braunii (Spenn.)
Fée
Aspidium viviparum Mett.
Polystichum heterolepis Fée
(ambig.)
Polystichum illyricum Hahne
Polystichum × illyricum Hahne
(ambig.)
Polystichum lonchitis (L.) Roth Polystichum lonchitis (L.) Roth
P. aculeatum (L.) Roth
Strahinjčica Mt 1973
P. aculeatum (L.) Roth
P. aculeatum (L.) Roth
P. aculeatum (L.) Roth
P. aculeatum (L.) Roth
Visibaba 1975
NP Učka 1973
Kašina 1993
Sunger 2010
P. braunii (Spennet) Fée
Aspidium viviparum Fée
yes
1904
Strahinjčica Mt 1972
P. lonchitis (L.) Roth
P. lonchitis (L.) Roth
Kalnik Mt 1961
Medak 1963
Treskavica (Bosnia &
Herc.) 1962
Badanj, Dinara Mt 1964
Prokletije (Montenegro)
1973
NP Velebit 1975
P. lonchitis (L.) Roth
P. lonchitis (L.) Roth
P. lonchitis (L.) Roth
val. P. aculeatum (L.) Roth
(acc. to FCD)
yes
P. illyricum Borbás
P. lonchitis (L.) Roth
unknown taxon, probably
culton
reconstructed card, illegible
source & year
syn. Dryopteris vivipara (Fée)
Kuntze
val. P. x illyricum (Borbás)
Hahne (acc. to FCD)
188 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
P. lonchitis (L.) Roth
P. lonchitis (L.) Roth
Polystichum munitum (Kaulf.)
C.Presl
Polystichum polyblepharum
(Roem. ex Kunze) C. Presl
Polystichum proliferum (R.Br.)
C.Presl
Polystichum setiferum (Forssk.)
T.Moore ex Woynar
Polystichum munitum (Kaulf.)
C. Presl
Polystichum polyblepharum
(Roem. ex Kunze) C. Presl
Aspidium proliferum R. Br.
(ambig.)
Polystichum setiferum (Forssk.)
Moore ex Woyn.
Aspidium munitum Kaulf.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
NP Velebit 1983
Komna (Slovenia) 1985
1904
?
card is missing
yes
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Aspidium proliferum Br.
Polystichum angulare
1904
P. setiferum (Forsskål) Woynar
Bosiljevo 2008
P. setiferum (Forsskål) Woynar
Strahinjčica Mt 1979
Polystichum setiferum (Forssk.) Polystichum setiferum (Forssk.) P. setiferum 'Plumosum
T.Moore ex Woynar (cult.)
Moore ex Woyn. (cult.)
Densum'
Buzet Nursery 2006
Polystichum (angulare?)
proliferum Wollastoni
1904
val. P. setiferum (Forssk.)
Woyn. (acc. to FCD)
val. P. setiferum (Forssk.)
Woyn. (acc. to FCD)
P. setiferum (Forssk.) Moore ex
Woyn. 'Plumosum Densum'
P. setiferum (Forssk.) Moore ex
Woyn. 'Proliferum
Wollastonii'
Order/clade & Family: POLYPODIALES – Lonchitiaceae (PPG-1, 2016 via APweb, 2021)
Lonchitis hirsuta L.
unknown
Szeged (Hungary) 1954
Pteris laciniata Willd.
in hand-written inventory of
glasshouse
Order/clade & Family: POLYPODIALES – Nephrolepidaceae (PPG-1, 2016 via APweb, 2021)
Nephrolepis Schott
(Polypodiaceae)
Nephrolepis
?
Nephrolepis biserrata (Sw.)
Schott
Nephrolepis biserrata (Sw.)
Schott
N. biserrata Sw.
N. rufescens
yes
Cambridge (UK) 1954
reconstructed card
Coimbra (Spain) 1955
in hand-written inventory of
glasshouse in 1955
191?
1932
189
N. biserrata Schott.
Not existing at the time
(Heinz, 1895-6)? Perhaps
under some other name
?
Tab. 1. Continued
PoWO (Kew):
WFO:
Nephrolepis biserrata (Sw.)
Schott (cult.)
Nephrolepis bostoniensis Anon.
(unplaced)
Nephrolepis biserrata (Sw.)
Schott (cult.)
Nephrolepis bostoniensis
(ambig.)
unknown
unknown
Nephrolepis cordifolia (L.) C.
Presl
Nephrolepis cordifolia (L.) C.
Presl var. cordifolia
Nephrolepis cordifolia (L.) C.
N. cordifolia (L.) Presl
Presl
Nephrolepis imbricata C.Presl
Nephrolepis imbricata
(ambig.)
Nephrolepis cordifolia (L.) C.
Presl (cult.)
Nephrolepis cordifolia (L.) C.
Presl (cult.)
1904
– 1948
N. biserrata (Sw.) Schott
‘Furcans’
Nephrolepis Bostoniensis
yes
191?
1932
Sofia (Bulgaria) 1982
yes
yes
gift 2016
Probably a cultivar of N.
exaltata (Bostoniensis')
reconstructed card; probably
cult. of N. exaltata group
new sprouted with others
during growing from spores
in hand-written inventory of
glasshouse
unknown cultivar; perhaps of
N. cordifolia 'Duffii' group; the
"Lemon Button Fern"
1904
191?
yes
Nephrolepis exaltata (L.) Schott
N. exaltata Schott var. selecta
(cult.)
Nephrolepis exaltata (L.) Schott
N. exaltata hort.
(cult.)
yes
Also in all inventories during
1950-ies. Planted outdoors in
2018
yes
N. exaltata (L.) Schott ‘Selecta’
1958
Nephrolepis exaltata (L.) Schott Nephrolepis exaltata (L.) Schott
N. exaltata (L.) Schott (cult.)
(cult.)
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
2016
1955
N. 'Pearls of Living' (?)
N. exaltata Schott
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
N. cordata Presl 'Compacta'
Nephrolepis exaltata (L.) Schott Nephrolepis exaltata (L.) Schott N. exaltata (Schott)
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
Heinz,
1895-6
2016
new sprouted with others
during growing from spores,
aff. 'Curly Locks'
N. exaltata crispa
1904
Nephrolepis exaltata 'Crispa'
N. "sesqu.filium." (?)
1904
illegible handwriting
N. exaltata 'Teddy'
Opeka Castle 1963
N. exaltata 'Teddy Junior'
Lada Nursery (Slovenia)
reconstructed card
1964
reconstructed card
190 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Nephrolepis hirsutula (G. Forst.)
C. Presl
Nephrolepis muscosa Clute
(unplaced name)
Nephrolepis hirsutula (G.
Forst.) C. Presl
Nephrolepis muscosa Clute
(ambig.)
Heinz,
1895-6
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
N. hirsutula (G. Forst.) Presl
yes
N. muscosa Pierson
p.p.
Szeged (Hungary) 1955
N. muscosa
Nephrolepis piersonii Anon.
(unplaced name)
Nephrolepis exaltata (L.) Schott N. exaltata Schott var. piersonii
(cult.)
F.R.Pierson
yes
N. Piersonii
191?
1932
N. Piersonii
yes
N. Piersonii fragrantissima
191?
in hand-written inventory of
glasshouse
N. exaltata (L.) Schott
'Piersonii'
in hand-written inventories of
glasshouse in 1950-ies
Nephrolepis piersonii Anon.
(unplaced name)
Nephrolepis whitmani Anon.
(unplaced name)
Nephrolepis sp. (unknown)
Nephrolepis sp. (unknown)
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis exaltata (L.) Schott
(cult.)
Nephrolepis sp. (unknown)
Nephrolepis sp. (unknown)
N. exaltata Schott var.
whitmannii Barrows
Nephrolepis Sieboldii
Nephrolepis Barousi
Nephrolepis sp. (unknown)
Nephrolepis sp. (unknown)
Nephrolepis Witmanii
191?
1932
Nephrolepis sp. (unknown)
Nephrolepis sp. (unknown)
Nephrolepis "Reppei" or
"Zeppei" (illegible)
1904
unknown
Nephrolepis sp. (unknown)
Nephrolepis sp. (unknown)
Nephrolepis "Philippiensis"
1904
unknown culton from
horticultural literature (e.g.
Birkenhead, 1892)
N. exaltata (L.) Schott
'Whitmannii'
unknown Combination
unknown
old cultivar from the Garden
catalogues in 1902-ies; perhaps
misspelled N. whitmani
yes
1904
191?
1904
yes
1904
Vodnjan Nursery 2021
new plants purchased in 2020
FCD: Woodsiaceae
191
Order/clade & Family: POLYPODIALES – Onocleaceae (PPG-1, 2016 via APweb, 2021)
Onoclea sensibilis L.
Onoclea sensibilis L.
O. sensibilis
(Aspleniaceae)
O. sensibilis L.
Onoclea struthiopteris (L.) Roth Matteuccia struthiopteris (L.)
Onoclea struthiopteris
(Aspleniaceae)
Tod.
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Matteuccia struthiopteris (L.)
Tod.
M. struthiopteris (L.) Tod.
M. struthiopteris (L.) Tod.
M. struthiopteris (L.) Tod.
Order/clade & Family: POLYPODIALES – Polypodiaceae (PPG-1, 2016 via APweb, 2021)
Campyloneurum crassifolium
(L.) Christenh.
Niphidium crassifolium (L.)
Lellinger
Ctenitis sloanei (Poepp. ex
Spreng.) C.V.Morton
Drynaria drynarioides (Hook.)
Christenh.
Drynaria meyeniana (Schott)
Christenh.
Aspidium furcatum Klotzsch
(ambig.) (Tectariaceae)
Aglaomorpha drynarioides
(Hook.) M.C. Roos
Aglaomorpha meyeniana
Schott
Polypodium morbillosum C.
Presl (ambig.)
Drynaria quercifolia (L.) J. Sm.
Buzet Nursery 2006
Vukova Gorica 2008
Macelj Mt 2012
Pleopeltis crassifolium (L.)
Moore/P. crassifolia T.Moore/ =
Polypodium crassifolium L.
Aspidium furcatum
1904
Vienna-Belvedere
(Austria) 2016
Vienna-Belvedere
(Austria) 2016
Aglaomorpha meyeniana Schott
Polypodium morbilosum
D. quercifolia (L.) J.Sm.
Goniophlebium subauriculatum
(Blume) C.Presl
Two different authors, two
different valid names (one of
them is Goniophlebium
subauriculatum (Blume) C.
Presl)
Schellolepis subauriculata
(Blume) J. Sm.
Goniophlebium subauriculatum
(Blume) C.Presl
Oreogrammitis reinwardtii
(Blume) Parris
Polypodium Reinwartdi/i
Lepisorus longifolius (Blume)
Holttum
unknown (P. longifolium Cav.*) Polypodium longifolium Mett.
Polypodium longifolium
plant labelled as "Polypodium
polycarpon (P. punctatum)"
(no author)=val. Microsorum
punctatum (L.) Copel.
(p.p.)
Aglaomorpha drynarioides
(Hook.) Roos
Drynaria quercifolia (L.) J.Sm.
Four different authors and valid
(?)
synonyms
Arilja (Serbia) 1987
1904
*incorrect; Drynaria J. Smith =
Aglaomorpha Schott (APG)
yes
Vienna-Belvedere
(Austria) 2016
1904
191?
1932
(p.p.)
1932
1950-ies
*P. longifolium Cav. =
Niphidium longifolium (Cav.)
C.V. Morton & Lellinger
(WFO)
192 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Lepisorus platyrhynchos
(Kunze) Li Wang
Leptochilus pteropus (Blume)
Fraser-Jenk.
Belvisia platyrhynchos (Kunze)
Copel.
Microsorum pteropus (Blume)
Copel.
Microgramma piloselloides (L.)
Copel.
Microsorum cuspidatum (D.
Don) Tagawa
Microsorum musifolium Copel.
'Crocodyllus'
Microsorum punctatum (L.)
Copel.
Microgramma piloselloides (L.)
Copel.
Polypodium leiorhizum Wall.
(ambig.)
Microsorum musifolium Copel.
(ambig.) 'Crocodyllus'
unknown (P. punctatum
Thunb. = val. Cochlidium
punctatum (Raddi) L.E. Bishop)
Microsorum punctatum (L.)
Copel.
Heinz,
1895-6
1904
– 1948
Belvisia platyrhynchos (Kunze)
Copel.
Microsorium pteropos (no
author)
M. pteropus (no author)
M. pteropus (no author)
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
München (Germany)
2016
1964
1981
1982
Riga (Latvia) 2016
M. piloselloides (L.) Copel.
Polypodium leiorhizum Wall.
*incorrect: Pteris sp.
yes
Microsorum 'Crocodyllus'
private collector
(Germany) 2020
Polypodium punctatum Sw.
1955
Polypodium punctatum Sw.
(=P.p. (L.) Sw.); changed to P.
polycarpon Cav.
(p.p.)
Microsorium punctatum (L.)
Cop.; changed to Polypodium
punctatum (no author)
(p.p.)
Phlebodium areolatum (Willd.) Phlebodium pseudoaureum
Phlebodium 'Pseudoaureum'
J.Sm. (cult.)
(Cav.) Lellinger (cult.)
Phlebodium aureum (L.) J.Sm. Phlebodium aureum (L.) J. Sm. Polypodium aureum L.
Phlebodium aureum (L.) J. Sm. Polypodium aureum L. var.
(cult.?)
umbellatum
Platycerium alcicorne (P.
Willemet) Desv.
Platycerium
alcicorne Desv. (ambig.)
Platycerium alcicorne Desv.
1958
2016
yes
Polypodium polycarpon Cav.=
Microsor(i)um punctatum (L.)
Copel. (WFO); Ple(i)opeltis
crassifolium (L.) Moore = val.
Niphidium c. (L.) Lellinger)!
this plant was labelled as P.
longifolium Mett.= val.
Lepisorus longifolius (Blume)
Holttum (Kew)
sprouted among others; aff.
'Blue Star'
yes
yes
1904
in hand-written inventory of
glasshouse
Ph. aureum (L.) J. Sm.
'Umbellatum' (?) – unknown
(cult.)/var.; I cannot see the
difference between sp. and cv.
193
Phlebodium aureum (L.) J. Sm.
(cult.?)
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Platycerium alcicorne
Heinz,
1895-6
yes
P. alcicorne (Willemet) Desv.
"Platycerium bicorne" (probably
P. x 'Bicorne') – perhaps
"biforme"?
Platycerium sp. (unknown)
Platycerium sp. (unknown)
Platycerium bifurcatum (Cav.)
C.Chr.
Platycerium bifurcatum (Cav.)
Platycerium bifurcatum
C. Chr.
Platycerium hillii T.Moore
Platycerium hillii T.Moore
(ambig.)
Polypodium L.
Polypodium sp. (?)
Polypodium sp. (?)
Polypodium sp. (?)
Polypodium sp. (?)
Polypodium sp. (?)
Polypodium sp. (?)
Four different authors and four
different val. syn.
Polypodium sp. (?)
Polypodium sp. (?)
Lophosoria quadripinnata (J.F.
P. glaucum
Gmel.) C. Chr. (Dicksoniaceae)
P. vulgare L. ssp. serratum
Polypodium cambricum L.
Willd.
Polypodium cambricum L.
Polypodium argenteum
Polypodium grandifolium Mett.
(unknown author)
Polypodium sagilifolia
Polypodium pygnosurum
(p.p.)
(p.p.)
(p.p.)
1961
Zagreb Nursery 2009
in hand-written inventories of
glasshouse in 1950-ies; hybrid
of P. Bifurcatum x P. Alcicorne
I cannot see the difference; P.
alcicorne is often sold as "P.
bifurcatum of gardens"
1904
191?
1932
Platycerium Hillei
Polypodium spp.
191?
1932
yes
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
yes
Platycerium bifurcatum (Cav.)
Platycerium willinckii T.Moore
C.Chr. subsp. willinckii (T.
Platycerium Willinski
(ambig.)
Moore) Hennipman & M.C.Roos
Platycerium Willinkii
Platycerium willinckii T.Moore
Platycerium vassei Anon.
Platycerium
P.vassei; changed to P. sp.
(unplaced name)
alcicorne Desv. (ambig.)
Platycerium wallichii Hook.
Platycerium wallichii Hook.
Platycerium Wallichi
Platycerium wilhelminaePlatycerium wilhelminae
P. wilhelminae-reginae v.A.v.R.
reginae Alderw. (unplaced)
Reginae v.A.v.R. (ambig.)
Polypodium L.
1904
– 1948
1904
191?
Riga (Latvia) 2016
(p.p.)
Zagreb Nursery 2001
plant looks like P. alcicorne
191?
yes
yes
OK
1904
191?
1958
1904
191?
191?
Kukuljanovo 1968
probably Platycerium ‘Queen
Wilhelmina'
"by the Grotto" (also in
glasshouses)
Perhaps P. argenteum Jacq.?
Acc. Dietrich, 1837
in hand-written inventory of
glasshouse in 1958
unknown
Perhaps P. pycnosorus?
194 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
P. australe Fée
P. vulgare L. ssp. serratum
Willd.
P. vulgare L. ssp. serratum
Willd.
P. australe Fée
P. cambricum L.
Polypodium vulgare L.
Polypodium vulgare L.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Dubrovnik 1978
Lapad 1969
Pelješac 1971
Polače 1979
Dubrovnik 2001
P. vulgare L.
1917
P. vulgare L.
yes
Hand-written inventory of
Medicinal plants
in hand-written inventories of
Karstic rockery in 1950-ies
P. vulgare L.
P. vulgare L.
P. vulgare L.
P. vulgare L.
Klek 1961
Rovinj 1963
NP Medvednica 1969
Treskavica (Bosnia &
Herc.) 1962
Vinica 1969
Brezovica 1970
Samobor 1980
Vukova Gorica 2008
Cyclophorus abbreviatus C.Cr.
Antwerp (Belgium) 1965 reconstructed card
Cyclophorus abbreviatus C.Cr.
Tübingen (Germany)
1971, 1973, 1979
Frankfurt (Germany)
lost or overwritten card
1958
Leiden (the Netherlands)
reconstructed card
1985
P. vulgare L.
P. vulgare L.
P. vulgare L.
P. vulgare L.
Pyrrosia abbreviata (Zoll.)
Tagawa
Cyclophorus abbreviatus C.
Chr. (ambig.)
Pyrrosia adnascens (Sw.) Ching Pyrrosia lanceolata (L.) Farw.
Nipholobus adnascens Klf. /(Sw)
Kaulf./
Pyrrosia christii (Giesenh.)
P. christii (Giesenh.) Ching
Ching
Pyrrosia lingua (Thunb.) Farw. Polypodium lingua Sw.
Pyrrosia nummulariifolia (Sw.)
P. nummulariifolia (Sw.) Ching
Ching
(glasshouse)
1904
yes
195
Pyrrosia christii (Giesenh.)
Ching
Pyrrosia lingua (Thunb.) Farw.
Pyrrosia nummulariifolia (Sw.)
Ching
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
1904
– 1948
Polypodium glaucum (changed
Four different authors leading to Lophosoria quadripinnata (J.F.
to Phymatodes glaucum;
four genera!
Gmel.) C. Chr.
authors missing)
Serpocaulon triseriale (Sw.)
Polypodium pycnosorum Link
Polypodium "pycnorosum" (?)
A.R.Sm. (?)
(ambig.)
Order/clade & Family: POLYPODIALES – Pteridaceae (PPG-1, 2016 via APweb, 2021)
private collector
(Germany) 2021
Vienna-Belvedere
(Austria) 2016
Actiniopteris radiata (Sw.) Link
Adiantum L.
Adiantum sp.
Adiantum spp.
Adiantum sp. cult.
yes
1954
Vienna-Belvedere
(Austria) 2016
ELTE Budapest
(Hungary) 2017
Adiantum sp. cult.
Adiantum sp. cult.
Adiantum aethiopicum L.
Adiantum chilense Kaulf. var.
sulphureum (Kaulf.) Kuntze ex Adiantum aethiopicum
Hicken
191?
Adiantum bausei T.Moore
(unplaced)
Adiantum bausei T.Moore
(ambig.)
Adiantum Bausei
1904
191?
Adiantum williamsii Moore
(ambig.)
Adiantum Wiliamsii
1904
Adiantum chilense Kaulf. var.
sulphureum (Kaulf.) Kuntze ex
Hicken
Adiantum capillus-veneris L.
Adiantum capillus-veneris L.
Adiantum capillus-veneris
Adiantum capillus Veneris
A. capillus Veneris
A. capillus-veneris L.
Probably misspelled
1904
Actiniopteris radiata (Sw.) Link Actiniopteris radiata (Sw.) Link Actiniopteris radiata
Adiantum L.
Adiantum sp.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
card is lost; most probably
orig. Polypodium glaucum
Cluj-Napoca (Romania)
p.p.
(Brack) Kuntze (syn.
1968
Phymatodes glauca (Brack.) J.
Sm.)
yes
yes
"various"; FCD: Adiantaceae
reconstructed card
various, among others
young plantlets (gift)
"Gartenwelt" from 1904 names
this taxon as hyb. of A.
trapeziforme x A. decorum
(val. A: raddianum)
191?
1932
yes
2016
in hand-written inventories of
glasshouse during 1950-ies
among others
196 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Adiantum capillus-veneris L.
(cult.)
Adiantum caudatum L.
Adiantum concinnum Humb. &
Bonpl. ex Willd.
Adiantum capillus-veneris L.
(cult.)
Adiantum caudatum L.
Adiantum concinnum Humb.
& Bonpl. ex Willd.
Adiantum diaphanum Blume
Adiantum diaphanum Blume
Heinz,
1895-6
1904
– 1948
A. capillus-veneris L.
A. raddianum
Adiantum concinum
Adianthum concinum
Adiantum diaphanum Blume
Adiantum Diaphanum
1904
191?
1932
1904
191?
Vienna-Belvedere
(Austria) 2016
A. diaphanum Blume
Adiantum edgeworthii Hook.
Adiantum edgeworthii Hook.
Adiantum elegantissimum
Anon. (unplaced)
Adiantum gracillimum T.Moore
(unplaced)
Adiantum elegantissimum ht.
(ambig.)
Adiantum gracillimum T.
Moore (ambig.)
Adiantum Edgevvoiti
Adiantum Egeworthii
Adiantum elegantissima/um
Adiantum hispidulum Sw.
Adiantum hispidulum Sw.
1904
191?
1904
191?
Adiantum Gracilimum/
gracillinum
1904
A. gracilimum
191?
1932
Adiantum groenewegenianum Adiantum groenewegenianum
Adiantum graenevegenianum
Regel (unplaced)
Regel (ambig.)
A. groenevegenianum
1904
191?
Vienna-Belvedere
(Austria) 2016
Riga (Latvia) 2016
Vienna-Belvedere
(Austria) 2016
A. hispidulum Sw.
A. hispidulum Sw.
Adiantum macrophyllum Sw.
Adiantum macrophyllum Sw.
A. macrophyllum Sw.
Adiantum monochlamys
D.C.Eaton
Adiantum monochlamys
D.C.Eaton
Adiantum Veitchii
Adiantum moorei Baker
Adiantum Veitchi
Adiantum raddianum C. Presl Adiantum "Morellii"
1904
191?
1904
191?
1932
Adiantum veitchii Hance
197
Adiantum "moorelli"
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
among others, aff. 'Yellow
(several different) 2016
Petticoat'
Zagreb Nursery 2012
*incorrect (A. caudatum L.)
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Adiantum legrandii Veitch
(unplaced)
Adiantum patens Willd.
Adiantum legrandii ht. Veitch
(ambig.)
Adiantum patens Willd.
Heinz,
1895-6
Adiantum legrande
1904
Adiantum patens Willd.
1904
191?
1932
A. patens
Adiantum pedatum L.
Adiantum pedatum L.
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Cluj-Napoca (Romania)
1969
Oslo (Norway) 1980
*incorrect
Stockholm (Sweden)
1982
A. pedatum L.
A. pedatum L.
A. pedatum L.
Adiantum peruvianum
Klotzsch
Adiantum peruvianum
Klotzsch
Adiantum peruvianum
Klotzsch
1904
Adiantum polyphyllum Willd.
Adiantum grande (ambig.)
Adiantum grande
1904
191?
1932
three different authors (none of
which is Schkuhr), with
Adiantum pubescens Schkuhr Adiantum pubescens
different valid names!
Adiantum raddianum C. Presl Adiantum raddianum C. Presl Adiantum cuneatum Langd.
A. Cuneatum
Adiantum raddianum C. Presl
(cult.)
Adiantum raddianum C. Presl
(cult.)
Adiantum raddianum C. Presl
(cult.)
Adiantum raddianum C. Presl
(cult.)
Adiantum raddianum C. Presl
(cult.)
Adiantum raddianum C. Presl
(cult.)
Adiantum raddianum C. Presl
(cult.)
Adiantum raddianum C. Presl
(cult.)
1904
yes
1904
fam. Polypodiaceae; in
hand-written inventories in
1950-ies
yes
A. raddianum C. Presl
yes
A. brillantelse
yes
A. cuneatum var. gracillimum
yes
A. decorum Moore
‘Magnificum’
A. raddianum Presl
'Microphyllum'
Vienna-Belvedere
(Austria) 2016
A. raddianum C. Presl
‘Brilliantelse’
A. raddianum C. Presl
‘Gracillimum’
A. decorum Moore
‘Magnificum’
yes
Vienna-Belvedere
(Austria) 2016
198 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
PoWO (Kew):
WFO:
Adiantum raddianum C. Presl
(cult.)
Adiantum raddianum C. Presl
Adiantum cuneatum fol. var.
(cult.)
Heinz,
1895-6
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
1904
191?
Vienna-Belvedere
(Austria) 2016
A. raddianum Presl 'Variegatum'
Adiantum rhodophyllum T.
Moore (unplaced)
Adiantum rhodophyllum T.
Adiantum rhodophyllum
Moore (ambig.)
Adiantum farleyense T.Moore
Adiantum tenerum Sw.
Adiantum Farleyense
(ambig.)
Adiantum fergusonii T.Moore
Adiantum Fergusonti
(ambig.)
Adiantum Fergusonthi
A. tenerum var. scutatum hort f.
Adiantum tenerum Sw. (cult.)? Adiantum tenerum Sw. (cult.)?
roseum
Adiantum tetraphyllum Humb. Adiantum tetraphyllum Humb.
A. tetraphyllum Willd. *
& Bonpl. ex Willd.
& Bonpl. ex Willd.
Adiantum trapeziforme L.
Adiantum trapeziforme L.
Adiantum trapeziforme L.
Adiantum trapesiforme
1904
yes
1904
191?
Anogramma leptophylla (L.)
Link
Ceratopteris thalictroides (L.)
Brongn.
191?
yes
yes
yes
Pityrogramma argentea
(Willd.) Domin
Gymnogramme aurea
Cryptogramma
acrostichoides R. Br.
Cryptogramma
acrostichoides R.Br.
Vienna-Belvedere
(Austria) 2016
NP Mljet 2016
yes
withdrew in 2018
1904
191?
C. thalictroides (L.) Brongn.
Cerosora argentea (Willd.)
Hennequin & H.Schneid. var.
aurea (Willd.) Hennequin &
H.Schneid.
Cryptogramma
acrostichoides R. Br.
*incorrect (A. capillus-veneris)
yes
1904
A. leptophylla (L.) Link
Ceratopteris thalictroides
A. tenerum Sw. 'Farleyense'?
1904
A. trapeziforme L.
Anogramma leptophylla (L.)
Link
Ceratopteris thalictroides (L.)
Brongn.
also sprouted among others
Nat. Croat. Vol. 31(1), 2022
Original name in the Garden
Database:
1961
val. Name for Gymnogramma
aurea (Willd.) Desv., fam.
Polypodiaceae
191?
Chemnitz (Germany)
2016
199
Tab. 1. Continued
WFO:
Hemionitis L. (p.p.)
Hemionitis L. (p.p.)
Gymnogramme spp.
Hemionitis achariorum
Christenh.
Doryopteris pedata (L.) Fée var.
Pteris palmata Willd.
palmata (Willd.) Hicken
Doryopteris palmata 'Pedata'
Hemionitis atropurpurea (L.)
Christenh.
Hemionitis calomelanos (Sw.)
Christenh.
Hemionitis lanosa (Michx.)
Christenh.
Heinz,
1895-6
1904
– 1948
yes
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Gymnogramme =
Gymnogramma Desv. fam.
Polypodiaceae
1904
Zagreb Nursery 2017
Pellaea atropurpurea (L.) Link P. atropurpurea (L.) Link
yes
Pellaea calomelanos (Sw.) Link Pellaea hastata Prantl.
no
1958
Cheilanthes lanosa (Michx.)
D.C. Eaton
Cheilanthes lanosa
Zagreb Nursery 2016
Zagreb Nursery 2020
Hemionitis marantae (L.)
Christenh.
Paraceterach marantae (L.)
R.M. Tryon
Hemionitis palmata L.
Hemionitis palmata L.
Cheilanthes lanosa
Notholaena marantae (L.) R.Br.
(changed to Cheilanthes
marantae (L.) Domin)
Hemionitis palmata
Kopaonik Mt (Serbia)
1963
in hand-written inventory of
glasshouse
Myriopteris lanosa
(Michx.) Grusz & Windham
(Serpentine species)
1932
Prague (Czech Republic)
reconstructed card
1964
Vienna-Belvedere
(Austria) 2016
Hemionitis rotundifolia (G.
Forst.) Christenh.
Hemionitis seticaulis (Hook.)
ined.
Hemionitis viridis (Forssk.)
Christenh.
Allosorus rotundifolius Kunze
Allosurus rotundifolia
(ambig.)
Pellaea rotundifolia (G.Forst.)
Pellaea rotundifolia Hook.
Hook
1904
yes
yes
Vienna-Belvedere
(Austria) 2016
Pellaea falcata Fée
Pellaea falcata (R.Br.) Fée
Riga (Latvia) 2016
Pellaea viridis (Forssk.) Prantl
Pellaea viridis (Fors.) Prantl
Vienna-Belvedere
(Austria) 2016
Bolbitis repanda (Blume) Schott.
unknown
(Polypodiaceae)
Gymnopteris repanda
Hamburg 1958
in a hand-written inventory of
greenhouse in 1958.
"Gymnopteris" could be a
synonym of Hemionitis or
Leptochilus
200 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Llavea cordifolia Lag.
Llavea cordifolia Lag.
Llavea cordifolia Lag.
Heinz,
1895-6
Gymnogramme calomelanos
Pityrogramma calomelanos (L.) Pityrogramma calomelanos (L.)
(Kaulf.); changed to Ceratopteris
Link
Link
calomelanos Und.
Gymnogramma massonii
Gymnogramme massonium
Loud. (ambig.)
Pityrogramma chrysoconia
Gymnogramma decomposita
Gymnogramme decomposistum
Maxon ex Domin
Baker (ambig.)
Pityrogramma chrysophylla
Gymnogramma
Gymnogramme chrysophylla
Link var. chrysophylla
chrysophyllum Kaulf. (ambig.)
Pityrogramma schizophylla
Pityrogramma schizophylla
(Baker ex Jenman) Maxon
(Baker) Maxon (syn.
(perhaps Gymnogramma
Gymnogramme gloriosa
Gymnogramma schizophylla
gloriosa Hort., acc. to GBIF,
Baker) (cult.)
2021)
Gymnogramma laucheana
Gymnogramme Laucheana
K.Koch (ambig.)*
Pityrogramma sulphurea (Sw.) Pityrogramma sulphurea (Sw.)
Maxon
Maxon
1904
– 1948
1904
191?
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Vienna-Belvedere
(Austria) 2016
yes
1904
Probably this species
1904
Probably this species
1904
191?
1904
In horticultural magazines of
19the century as
Gymnogramma
schizophyllum var. gloriosa
191?
*also Gymnogramma
laucheana hort.
yes
191?
1932
Gymnogramme sulfurea
Gymnogramme sulphurea Desv.
Pteris L.
Pteris spp.
Pteris argyraea T.Moore
Pteris argyraea T.Moore
(ambig.)
P. argyraea
Pteris sp.
Pteris sp.
Pteris sp.
Pteris sp.
P. argyraea Moore
P. argyraea crispa
P. argyraea v. grandis
Berlin (Germany) 1955
yes
1904
191?
1932
191?
191?
in hand-written inventory of
glasshouse in 1955
"By the Grotto", in the
glasshouses; "fam.
Polypoidaceae"
yes
Perhaps Pteris crispa?
Perhaps Pteris grandis?
201
Pteris L.
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
(four P. arguta with different
authors)
Pteris biaurita L.
Pteris cretica L.
(three P. arguta with different
authors)
Pteris biaurita L.
Pteris cretica L.
Heinz,
1895-6
1904
– 1948
yes
1932
"Pteris arguta"
P. biaurita L.
Pteris cretica
P. cretica L.
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
in hand-written inventory of
1951
glasshouse
yes
fam. Polypodiaceae
new sprouted among others
yes
2016
grown from spores
1904
191?
Pteris cretica L. (cult.)
Pteris cretica L. (cult.)
P. cretica v. albolineata
Pteris cretica L. (cult.)
Pteris cretica L. (cult.)
P. cretica var. albolineata Hook.
Pteris cretica L. (cult.)
Pteris cretica L. (cult.)
P. cretica L. 'Albo-Lineata'
Pteris cretica L. (cult.)
Pteris cretica L. (cult.)
P. cretica L. var. cristata hort.
Pteris cretica L. (cult.)
Pteris cretica L. (cult.)
P. cretica major
Pteris cretica L. (cult.)
Pteris gauthieri hort. (ambig.)
P. cretica L. var. gauthieri hort.
yes
Pteris cretica L. (cult.)
Pteris cretica L. (cult.)
P. cretica L. var. roweri hort.
yes
Pteris cretica L. (cult.)
Pteris wimsettii ht. (ambig.)
P. cretica L. var. wimsettii hort.
yes
Pteris wimsettii ht. (ambig.)
P. Wimsetti
Pteris ensiformis Burm.f.
Pteris ensiformis Burm.f.
Pteris ensiformis Burm.
Pteris ensiformis
1904
191?
1904
191?
Pteris ensiformis Burm.f. var.
ensiformis
Pteris victoriae ht. (ambig.)
Pteris Victoria*
1904
*Probably Pteris victoriae
(W.Bull ex Ridl.) W. Bull
Pteris longifolia L.
Pteris longifolia L.
Pteris longifolia/us
1904
191?
1932
fam. Polypodiaceae
yes
yes
yes
2016
yes
2016
191?
1932
yes
in hand-written inventories of
glasshouse in 1950-ies
Hort: P. cretica L. 'Albolineata';
new sprouted among others
Hort: P. cretica L. 'Cristata';
new sprouted among others
grown from spores
probably P. cretica 'Major'
Hort: P. cretica L. 'Gauthieri'
Hort: Pteris cretica L.
‘Roweri’/'Roeweri'
Hort.: Pteris cretica L.
‘Wimsetti’/Wimsettii’
202 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
Pteris longifolia L.
1904
– 1948
yes
"=P. longifolia L., acc. to
Ehrendorfer"
Pteris multifida Poir.
unknown (Pteris serrulata
Forssk. – ambig.)
Pteris serrulata L.F.
P. serrulata
Pteris cristata Anon. (unplaced) Pteris cristata hort. (ambig)
Pteris multifida Poir. ‘Cristata’
yes
191?
yes
P. serrulata
yes
Pteris serrulata var. cristata
Pteris sp.
1904
P. multifida
P. serrulata L.f.; changed to P.
multifida Poir.
Pteris serrulata cristata
Pteris sp.
Pteris serrulata grandis L.f.
unknown (only Pteris serrulata
Pteris multifida Poir. ‘Cristata’
Forssk. – ambig.)
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
in hand-written inventories of
glasshouse in 1950-ies
In our database P. longifolia
was once considered to be a
yes
synonim of P. vittata (Kovačić,
2015)
Szeged (Hungary) 1954
(without an author, but we can
presume it was L.f. as in 1904;
fam. Polypodiaceae)
in hand-written inventories of
glasshouse in 1950-ies
in hand-written inventories of
glasshouse in 1950-ies
yes
1904
in hand-written inventories of
glasshouse in 1950-ies
perhaps Pteris grandis?
yes
1904
yes
Pteris quadriaurita Retz.
Pteris quadriaurita Retz.
P. quadriaurita Retz.
yes
Pteris vittata L.
Pteris vittata L.
P. vittata L.
yes
191?
1932
Vienna-Belvedere
(Austria) 2016
err. in Kovačić, 2015:
"Histiopteris incisa (Thunb.) J.
Sm. – arrived as syn. Pteris
quadriaurita (Pteridaceae)"
new sprouted among others
grown from spores; spreading
locally
203
Order/clade & Family: POLYPODIALES – Tectariaceae (PPG-1, 2016 via APweb, 2021)
Hypoderris brownii J.Sm.
Hypoderris brownii J. Sm. ex
Hypoderis browni
(Polypodiaceae)
Hook.
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Tectaria aspidioides Copel.
(Polypodiaceae)
Polypodium philippinum
Hook. (ambig.)
Polypodium Phillipense
Tectaria aurita (Sw.) S.Chandra
Stenosemia aurita (Sw.) C. Presl Polybotrya aurita
(Polypodiaceae)
Tectaria cicutaria (L.) Copel.
Tectaria cicutaria (L.) Copel.
Tectaria cicutaria (L.) Copel.
(Polypodiaceae)
two different authors and two Tectaria mexicana (Fée) C.V.
Aspidium latifolium
diffrerent species of Tectaria
Morton
Tectaria zeilanica (Houtt.)
Tectaria zeylanica (Houtt.)
Quercifilix zeylanica (Houtt.)
Sledge (Polypodiaceae)
Sledge
Copel.
Order/clade & Family: POLYPODIALES – Thelypteridaceae (PPG-1, 2016 via APweb, 2021)
Thelypteris beddomei (Baker)
Parathelypteris beddomei
Parathelypteris beddomei
Ching (Aspleniaceae)
(Baker) Ching
(Baker) Ching
Thelypteris balbisii (Spreng.)
Thelypteris balbisii (Spreng.)
Aspidium Balbisii
Ching (Aspleniaceae)
Ching
Thelypteris cordata (Fée)
Thelypteris cordata (Fée)
Th. cordata (Fée) Proctor
Proctor (Aspleniaceae)
Proctor
Thelypteris kunthii (Desv.)
Thelypteris kunthii (Desv.)
Th. kunthii (Desv.) C.V.Morton
C.V.Morton (Aspleniaceae)
C.V.Morton
Thelypteris noveboracensis (L.) Thelypteris noveboracensis (L.)
Aspidium novaeboracense
Nieuwl. (Aspleniaceae)
Nieuwl.
unknown (Nephrodium/
Thelypteris palustris Schott
Dryopteris thelypteris = val.
Dryopteris thelypteris A.Gray
(Aspleniaceae)
Thelypteris confluens (Thunb.)
C.V.Morton)
Nephrodium thelypteris (L.)
Strempel – Dryopteridaceae
*Phegopteris "cunata" Mett.
unknown
unknown
(misspelled?)
Order/clade & Family: POLYPODIALES – Woodsiaceae (PPG-1, 2016 via APweb, 2021)
Woodsia obtusa (Spreng.) Torr.
Woodsia obtusa Torr.
W. obtusa Torr.
(Aspleniaceae)
Heinz,
1895-6
1904
– 1948
191?
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
I am not sure that this is the
valid name(s) of the original
taxon
1904
yes
1904
Vienna-Belvedere
(Austria) 2016
Chemnitz (Germany)
2016
1904
private collector
(Germany) 2020
Antwerp (Belgium) 2016
1904
yes
1955
1962
Chemnitz (Germany)
2016
hand-written inventories of
Alpinum rockery in 1950-ies
Thelypteris palustris Schott
(FCD)
perhaps Ph.cruciata (Willd.)
Mett.?
204 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
PoWO (Kew):
Original name in the Garden
Database:
Tab. 1. Continued
WFO:
Heinz,
1895-6
Woodsia plummerae Lemmon
Woodsia plummerae Lemmon W. plummerae Lemmon
(Aspleniaceae)
Woodsia scopulina D.C. Eaton
Woodsia scopulina D.C. Eaton Woodsia scopulina subsp.
subsp. laurentiana Windham
(unknown subsp.)
laurentiana
(Aspleniaceae)
Order/clade & Family: various, undeterminated
Aspidium spp. (p.p. synonym of Aspidium spp. (synonym of
Tectaria Cav.; Tectariaceae)
Tectaria Cav.; Polypodiaceae)
Aspidium spp.
1904
– 1948
Source (in Croatia, if not
Kovačić, stated otherwise) and
Comment
2015 year of obtainig. Empty
box = unknown
Chemnitz (Germany)
2016
Chemnitz (Germany)
2016
yes
various
Nat. Croat. Vol. 31(1), 2022
PoWO (Kew):
Original name in the Garden
Database:
probably Croatian native, as
they grew "by the Grotto", but
some perhaps also of Tectaria
genus; today "Aspidium"-s are
scattered to various families
Tab. 1. Ferns inventoried in Botanical Garden of the Faculty of Science, University of Zagreb, between 1895 and 2021, including “hidden” entries retrieved
from the reused paper-cards and hand-written lists. Column 1 “PoWO (Kew)” depicts the views of each fern-name (December, 2021) as seen in the Plants of
the World Online, compared to the “WFO” (2nd column), showing the same name according to the World Flora Online, and (3rd column) “Original name in
the Garden Database”.
Columns 4, 5 and 6 contain the original sources of information for each fern: published in Heinz (1895-6) and/or Kovačić (2015), or unpublished (hand-written
inventories between 1904 and 1948). Column 7 carries the information on the source of plant material and year of its obtaining (if such exist). Last column
depicts additional data on each fern (for Croatian native species, Flora Croatica Database was consulted). Affiliations to Orders/Clades and Families are arranged in rows separating groups of ferns, according to The Pterydophyte Phylogeny Group (2016; via APweb, 2021).
205
206 Kovačić, S.: Plethora of Plants 7 – Collections of the Botanical Garden, Faculty of Science, University of Zagreb
1
2
3
4
Four seasons in the Garden fernarium: majority of species are planted outdoors, while tropical ones are
brought out with pots in May, to over-summer. 1) Spring. 2) Summer. 3) Autumn. 4) Winter.