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New distribution and taxonomic information on Callitriche (Plantaginaceae)

in the Mediterranean region

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DOI: 10.11646/phytotaxa.313.1.6

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 Phytotaxa 313 (1): 091–104 ISSN 1179-3155 (print edition)
http://www.mapress.com/j/pt/
Article PHYTOTAXA
Copyright © 2017 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.313.1.6

New distribution and taxonomic information on Callitriche (Plantaginaceae) in the

Mediterranean region
RICHARD V. LANSDOWN1*, IOANNIS BAZOS2, MARIA CARMELA CARIA3, ANGELO TROIA4 & JAN J.
WIERINGA5
1
45 The Bridle, Stroud, Glos. UK GL5 4SQ; e-mail: rlansdown@ardeola.demon.co.uk.
2
Institute of Systematic Botany, Department of Biology, School of Science, National and Kapodistrian University of Athens, Panepistimi-
opolis, 15784 Athens, Greece; e-mail: ibazos@biol.uoa.gr
3
Dipartimento di Scienze della Natura e del Territorio, Università degli Studi di Sassari, Sassari, Italy; e-mail: mccaria@uniss.it
4
Dipartimento STEBICEF, sezione di Botanica ed Ecologia Vegetale, Università degli Studi di Palermo, Palermo, Italy;
e-mail: angelo.troia@unipa.it
5
Naturalis Biodiversity Center, National Herbarium of The Netherlands, Darwinweg 2, 2333 CR Leiden, The Netherlands;
e-mail: jan.wieringa@naturalis.nl
* author for correspondence

Abstract

This article presents new records of water-starworts (Callitriche sp. pl.) from the Mediterranean basin, resulting from review
of herbarium specimens and field work. Callitriche brutia var. naftolskyi is stated as a new combination and confirmed from
Greece (Lesvos and Milos), Israel, Italy (Sardinia and Sicily), Libya, Morocco and Syria; C. lusitanica from Greece (Les-
vos), Israel and Italy (Sardinia and Sicily); C. brutia var. brutia has been known from Greece for some time but is confirmed
from Lesvos and Milos; C. obtusangula and C. truncata subsp. truncata are both confirmed from Sardinia and Sicily, while
the latter is also confirmed from Syria. Callitriche lenisulca and C. stagnalis are confirmed from Sardinia but records of the
latter from Lesvos appear to be erroneous; C. truncata subsp. occidentalis is known from Lesvos but records from Sardinia
and Sicily appear to be erroneous. Records of C. regis-jubae from Sardinia are probably misidentifications for C. brutia var.
naftolskyi. Records of C. brutia var. hamulata from the region are likely to be erroneous. A recent find of the alien C. ter-
restris in Spain is the first record for that country and the second record for Europe.

Key words: Callitriche brutia var. naftolskyi; Callitriche lusitanica; Callitriche terrestris; Greece; Sardinia; Sicily; Spain;
water-starworts

Introduction

Callitriche Linnaeus (1753: 969) (Plantaginaceae) is a sub-cosmopolitan plant genus occurring throughout temperate
regions of both hemispheres, including the peri-Antarctic islands but typically mainly at high altitude in the tropics
(Lansdown 2008). Approximately 75 species have been recognized worldwide (Lansdown 2008), of which 16 are
known to occur in the Mediterranean. Records of Callitriche include frequent and extensive misidentification due to
a combination of factors including a lack of taxonomic and nomenclatural clarification before the mid-20th century,
critical distinctions between similar species and the difficulty of locating and recognising identification features.
In recent years, clarification of the identification characters for Callitriche species occurring in the Mediterranean
basin (e.g. Schotsman 1967, 1973, 1977, 1982, Schotsman & Andreas 1974, Lansdown 2008) has enabled revision
of some herbarium specimens, as well as stimulated collection of new records, including review of the distribution
and taxonomy of members of the genus in Sardinia and Sicily in Italy, and the islands of Lesvos and Milos in Greece.
Information is presented here on new records and taxonomic clarification arising from this review, together with a new
report of C. terrestris Rafinesque (1808: 358) from Spain.

Accepted by Manuel B. Crespo: 6 Jun. 2017; published: 11 Jul. 2017 91

Material and methods

This article is based on analysis of relevant literature, revision of herbarium specimens and field work. Callitriche
specimens cited in this article have been reviewed from the following herbaria: ATHU, BM, BR, C, CGE, E, FI, G,
HUJ, K, L, P, RAB, SS, STU and U (abbreviations according to Thiers 2017). Specimens of C. brutia var. naftolskyi
have been deposited in SS, specimens of C. lusitanica from Lesvos in ATHU and of C. terrestris from Córdoba in
WAG. Plant names follow The Plant List (2013) apart from Isoetes, for which the names follow Troia et al. (2016),
and Callitriche.

Results and discussion

Callitriche brutia var. naftolskyi (Warburg & Eig) Lansdown comb. nov. Callitriche naftolskyi Warburg & Eig (1929:
84), basionym.
Lectotype (designated here; cited but not formally designated by Schotsman 1967):ISRAEL. Sharon Plain, north-east of Tel Aviv, 23 April
1927, Naftolsky No. 01853 (HUJ) [not seen].

Callitriche brutia var. naftolskyi has been overlooked, dismissed as a synonym of C. brutia Petagna (1787: 10) var.
brutia or treated as unresolved by most authors (e.g. Schotsman 1967, Schotsman 1977, Lansdown 2008). However,
a chance encounter by RVL with populations agreeing with the description of C. naftolskyi at a number of sites during
the International Symposium on Mediterranean Temporary Ponds held in Sassari in 2015, led to a re-evaluation of its
status which is here regarded at the varietal rank (see below).
Callitriche naftolskyi was described as a new species by Warburg & Eig (1929), based on material from the Sharon
Plain, NE of Tel Aviv in Israel (lectotype: Naftolsky 01853, HUJ; cf. Schotsman 1967). Subsequently, Schotsman (1967)
initially considered single specimens from Sardinia (Huet de Pavillon s.n. 1854, P) and Morocco (Nègre 4876, RAB) to
be C. naftolskyi, but then expressed doubt (Schotsman 1977) and ultimately assigned them to C. brutia ( C. brutia var.
brutia). She also annotated a specimen from Palermo (Birch s.n. 1854, K) “probably C. pedunculata De Candolle (in
Lamarck & Candolle 1805: 415) (= C. brutia var. brutia) with some big fruits and broad winged seeds”, but this material
clearly conforms to the original concept of C. naftolskyi. The discovery of populations in Sardinia prompted review
of material from the Isola di Capraia, formerly identified as C. regis-jubae Schotsman (1974: 25) (cf. Lansdown et
al. 2014) and this was also shown to agree with the description of C. naftolskyi. Fieldwork in April 2016 revealed C.
brutia var. naftolskyi to be fairly widespread in suitable habitat in Sardinia and present in at least two seasonal pool
complexes in Sicily. Furthermore, review of herbarium specimens also revealed specimens from Greece (Lesvos and
Milos), Libya, Morocco and Syria. This review of the literature and herbarium specimens suggests that C. brutia var.
naftolskyi is endemic to but widespread within the Mediterranean basin, occurring on larger islands and in North Africa
east to the Middle East.
The following description is based on a combination of biometrics from material collected at a range of sites in
Sardinia in April 2016, together with information from the protologue (Warburg & Eig 1929) and details provided by
Schotsman (1967): Stem much-branched, supported by water or when terrestrial, prostrate and creeping. Stem scales
with 10–19 cells, often irregular in outline; axillary scales fan-like with 6–8 cells. Floating leaves narrowly elliptic,
at most very shallowly notched at the apex, 1.3–2.4 mm wide × 2.5–4 mm long, petiole 0.3–2.3 mm, with 3–5 veins,
terrestrial leaves typically smaller. Bracts apparently lacking; flowers solitary in leaf axils, the most frequent pattern
involves a male flower opposed by a female; pollination appears to be through self-fertilisation; anther average 0.3–0.4
mm wide × 0.3 mm high; stigma 0.6–1 mm long, persistent, initially reflexed, then appressed to the side of the fruit;
peduncle 4–12 mm; fruit 0.9–1.6 mm wide × 1–1.5 mm high; wing 0.02–0.1 mm, radial thickenings in the wings
are very branched and interlinked; toward the base of the wing there are often polygonal cells aligned in radiating
directions, continuing the mesocarp cells.
Five Callitriche taxa in the Mediterranean region produce winged, pedunculate fruit. Of these, both C. mouterdei
Schotsman (1969: 617) and C. regis-jubae have persistent bracts approximately 1 mm long, long filaments (at least
post-anthesis), yellow pollen and erect or recurved styles. Callitriche brutia var. naftolskyi lacks bracts, the anther is
subsessile, pollen is colourless and the style is strongly reflexed and soon appressed to the side of the fruit. Callitriche
truncata Gussone (1826: 4) subsp. truncata differs from C. brutia var. naftolskyi in its leaves which are always single-

92 • Phytotaxa 313 (1) © 2017 Magnolia Press LANSDOWN ET AL.

veined, and lack of peltate scales on the stem and leaves. Callitriche brutia var. naftolskyi can be distinguished from
C. b. var. brutia by the wings on mature fruit which are wider and have an undulate margin; the fruit are matt (shiny
in var. brutia); the style is persistent and the fruit typically wider than long, although this can be difficult to measure
because of the shape of the wing. Although C. b. var. naftolskyi differs from C. b. var. brutia in a number of ways,
these are insignificant compared to the characters separating most species within the genus, and pending molecular
clarification it is appropriate to treat C. naftolskyi as a variety within C. brutia as C. brutia var. naftolskyi. It is of note
that the genome is the same size in both C. brutia var. brutia and C. b. var. naftolskyi (J. Prancl pers. comm.).
Callitriche brutia var. naftolskyi has not been found with subsessile fruit. All fruit are pedunculate including young,
green ones with peduncles 4 mm or more in length, even in aquatic material, and most fruit are on peduncles 10 mm or
more in length. This differs from C. brutia var. brutia which has subsessile fruit when growing in water, but pedunculate
fruit when terrestrialised (Lansdown 2008). Warburg & Eig (1929) suggested that the fruits of C. naftolskyi penetrate the
soil vertically which is considered by Schotsman (1967) unlike C. brutia and to more closely resemble C. deflexa A.
Braun ex Hegelmaier (1864: 58). However, this is not a very reliable difference given that the direction of entry into the
soil and even whether fruit grow under a mat of vegetation or penetrate into the soil, depends very much on the nature
of the substrate in which the plant is growing.
In Sardinia and Sicily C. brutia var. naftolskyi was recorded exclusively from temporary wetlands, both in well-
known species-rich sites such as the seasonal pool complexes at Giara di Gesturi and Monte Minerva in Sardinia and a
wetland complex near Buccheri and pools at Castello della Pietra in Sicily; but also in a relatively banal ditch running
through pasture east of Perdasdefogu in eastern Sardinia. In these sites it was recorded from a variety of habitats, all
of which are seasonally inundated, including deeply rutted areas on tracks, pools and hollows in pasture, as well as
open shallow hollows and a cupular pool on limestone “pavement”. In Sicily and Sardinia, C. brutia var. naftolskyi
was most frequently recorded with a range of species characteristic of seasonal pools such as Bellis annua Linnaeus
(1753: 887), Carex divisa Hudson (1762: 348), Crassula vaillantii (Willdenow 1798: 720) Roth (1827: 992), Eryngium
barrelieri Boissier (1844: 125), Juncus pygmaeus Richard ex Thuillier (1800: 178), Lythrum hyssopifolia Linnaeus
(1753: 447) and Ranunculus ophioglossifolius Villars (1789: 731); less often with Apium crassipes (W.D.J. Koch
ex Reichenbach 1825: tab. 21) Reichenbach (1863: 9), Callitriche brutia var. brutia, C. stagnalis Scopoli (1772:
251), Chamaemelum nobile (Linnaeus 1753: 894) Allioni (1785: 185), Corrigiola litoralis Linnaeus (1753: 271),
Damasonium bourgaei Cosson (1849: 47), Isoetes species (including I. histrix Bory 1844: 1167, I. longissima Bory
1844: 1165, and I. tiguliana Gennari 1861: 42), Lythrum borysthenicum (M.Bieb. ex Schrank 1822: 643) Litvinov
(1917: 209), L. thymifolia Linnaeus (1753: 447), Mentha pulegium Linnaeus (1753: 577), Myosotis sicula Gussone
(1843: 214), Pilularia minuta Durieu in Bory & Durieu (1848: plate 38, figs. 120), Ranunculus muricatus Linnaeus
(1753: 555) and R. revellieri Boreau (1857: 85). It was also recorded once with each of the following species varying
from what may be described as “true” aquatics such as Ranunculus aquatilis Linnaeus (1753: 556) and R. peltatus
Schrank (1789: 103), through species typical of seasonal pools such as Antinoria insularis Parlatore (1845: 93) and
Juncus heterophyllus Dufour (1825: 88), to those which rarely, if ever actually grow in water but are characteristic
of damp ground including Cicendia filiformis (Linnaeus 1753: 231) Delarbre (1795: 20), Juncus bufonius Linnaeus
(1753: 328), J. capitatus Weigel (1772: 28) and Pulicaria sicula Moris (1840: 363).
No assessment of the extinction risk of C. brutia var. naftolskyi has been published, the extent of its range,
combined with the occurrence of extensive populations in many different water bodies in Sardinia and Sicily mean that
it should be classed as Least Concern (IUCN 2012).

Callitriche lusitanica Schotsman (1961: 112)

In 2003, a Callitriche specimen was sent to RVL by IB from between Parakila and Agra on Lesvos. It was tentatively
identified as C. lusitanica but could not be confirmed as some critical elements were lacking. Further material was
collected from the same site by IB and S. Zervou on 10 May 2014 and confirmed as C. lusitanica by RVL in January
2015. Subsequently, in the course of fieldwork, C. lusitanica was found at four sites scattered through the mountainous
central and eastern parts of Sardinia. In all of the Sardinian sites it was associated with the main channel of medium-sized
to large rivers, either in the channel itself, where it formed large stands, in backwaters, on the margins or growing over
boulders. In deeper water, it occurred with Callitriche obtusangula Le Gall (1852: 202), filamentous green algae, Lemna
gibba Linnaeus (1753: 970), Myriophyllum spicatum Linnaeus (1753: 992), Nasturtium officinale Brown (1812: 110),
Potamogeton crispus Linnaeus (1753: 126), P. schweinfurthii Bennett (1901: 220), Ranunculus peltatus, R. penicillatus
(Dumortier 1863: 216) Babington subsp. pseudofluitans (Syme 1863: 20) Webster (1988: 20) var. vertumnus Cook
(1966: 160), R. trichophyllus Chaix ex Villars (1786: 335), Sparganium erectum Linnaeus (1753: 971), and Stuckenia
Callitriche (Plantaginaceae) Phytotaxa 313 (1) © 2017 Magnolia Press • 93
pectinata (Linnaeus 1753: 127) Börner (1912: 713). On the margins and in backwaters, it was recorded with Apium
nodiflorum (Linnaeus 1753: 251) Lagasca (1821: 101), Corrigiola littoralis, Cyperus fuscus Linnaeus (1753: 46), C.
longus Linnaeus (1753: 45), Fontinalis hypnoides Hartman (1843: 434), Scirpoides holoschoenus (Linnaeus 1753:
49) Soják (1972: 127), Juncus articulatus Linnaeus (1753: 327), J. pygmaeus, Mentha × villosa-nervata Opiz (1831:
60), M. pulegium, Montia fontana Linnaeus (1753: 87) var. chondrosperma (Fenzl in Ledebour 1843: 152) Walters
(1956: 4), Oenanthe crocata Linnaeus (1753: 254), Parentucellia viscosa Caruel (1885: 482), Ranunculus muricatus,
R. ophioglossifolius, R. peltatus, Rumex conglomeratus Murray (1770: 52), Schoenoplectus lacustris (Linnaeus 1753:
48) Palla (1888: 299), Typha angustifolia Linnaeus (1753: 971) and Veronica anagallis-aquatica Linnaeus (1753: 12).
A notable feature of populations in Sardinia not recorded in populations in the Iberian Peninsula is the development of
extensive, evidently persistent terrestrial plants, both on the margins where water-levels had dropped but particularly
growing over boulders in the channel where they may remain humid for months.
Callitriche lusitanica is fairly widespread in the western half of the Iberian Peninsula, it has been recorded from
Algeria, Morocco, Tunisia and there is a specimen collected by the late Avinoam Danin from Golan in Israel (BR). A
plant found in Siracusa Province, Sicily by AT was identified as this species from a photograph. It is clearly much more
widespread than previously thought and likely to have been under-recorded throughout much of the region. The range
of C. lusitanica can now be seen to extend from the Iberian Peninsula and Morocco east through Sardinia and Lesvos
to Israel, suggesting that this species should be sought elsewhere in North Africa, as well as possibly in the Balearic
Islands.
In 2010, an assessment of the extinction risk of European populations of C. lusitanica classed it as Near Threatened
(Bilz et al. 2011) on the basis that “This species is classed as Near Threatened as it has a restricted distribution area and
its freshwater habitat is degrading. Expected future demand of water in the southern European regions, will worsen the
situation for this species and it could quickly fall into a threatened category”. The discovery of additional populations
extending from Sardinia and Sicily east to Syria and Israel means that its distribution is no longer restricted and
although its habitat is threatened throughout its range, the likelihood of extinction must now be considered remote.
Callitriche lusitanica should therefore be classed as Least Concern (IUCN 2012).

Callitriche terrestris Rafinesque (1808: 358)

In May 2016 a population of C. terrestris was found growing on moist soil between cobbles in the street in the Jewish
quarter of Córdoba (southern Spain), where it was dominant with a few other species such as Poa annua Linnaeus
(1753: 68), Plantago major Linnaeus (1753: 112), Sagina apetala Arduino (1764: 22) and Polycarpon tetraphyllum
(Linnaeus 1753: 89) Linnaeus (1759: 881) In its native range, C. terrestris grows in seasonally wet hollows and damp
soil, in cultivated fields, rocky ground, paths, tracks and on sandy river bars (Fassett 1951). It has been recorded only
once previously in Europe, on the margin of an acid pool (pH = 5.6) at Clairefontaine in the Forêt des Yvelines, Seine
et Oise Department in central France, in 1930 (Schotsman 1967, Lansdown 2008), but there are no recent records from
this site. Callitriche terrestris is native to the United States and southwards through Central America to Brazil and
Ecuador (Fassett 1951). It is naturalised in Taiwan (Lansdown 2006a) and has been reported from Japan (Katsuyama
& Kawasumi 1999), although the latter report has not been critically confirmed. The record from Spain therefore
represents the second one for Europe for this species. Since the previous record is over 85 years old, the population in
Spain may be considered a new introduction into Europe. Introduction from the Americas, especially North America,
is a known source for alien species in Europe (van Kleunen et al. 2015). Terrestrial Callitriche species are increasingly
occurring as contaminants in ornamental plant pots in international trade (unpublished data) and this is one possible
explanation for the route by which this plant entered the country; alternatively it could have been brought in as seed
carried accidentally by tourists visiting the nearby Mezquita-Catedral de Córdoba.

94 • Phytotaxa 313 (1) © 2017 Magnolia Press LANSDOWN ET AL.

FIGURE 1. Callitriche brutia var. naftolskyi, showing the fruit wider than high and broadly winged, near Perdasdefogu, Sardinia, 23 April
2016. Photograph by Richard V. Lansdown.

FIGURE 2. Terrestrial Callitriche brutia var. naftolskyi, near Perdasdefogu, Sardinia, 23 April 2016. Photograph by Richard V.
Lansdown.

Callitriche (Plantaginaceae) Phytotaxa 313 (1) © 2017 Magnolia Press • 95

FIGURE 3. Terrestrial Callitriche brutia var. naftolskyi, Paùli Piccia, Sardinia, 23 April 2016. Photograph by Richard V. Lansdown.

FIGURE 4. Dense beds of Callitriche lusitanica with flowering Ranunculus peltatus in a bedrock and cobble-dominated channel, Ollastu
River, Sardinia, 23 April 2016. Photograph by Richard V. Lansdown.

96 • Phytotaxa 313 (1) © 2017 Magnolia Press LANSDOWN ET AL.

FIGURE 5. Terrestrial Callitriche lusitanica growing over a boulder in the channel of the Rio Picocca, Sardinia, 23 April 2016. Photograph
by Richard V. Lansdown.

FIGURE 6. Callitriche lusitanica in a lowland, silt-dominated channel, Fiume Temo, Sardinia, 24 April 2016. Photograph by Richard V.
Lansdown.

Callitriche (Plantaginaceae) Phytotaxa 313 (1) © 2017 Magnolia Press • 97

FIGURE 7. Callitriche terrestris growing between cobbles in the street in Córdoba, 4th May 2016. Photograph by J.J. Wieringa.

Final comments

The first records of Callitriche species on Lesvos were made by Candargy (1897, 1898) who also described C. aeolica
Candargy (1897: 157) as a new species from Chamaris as follows “Submersa, stomatibus pilisque stellatis carens,
flores bibracteati, folia uninervia, linearia apice bifida, styli perlongi reflexo patentes, ovarium breviter pedicellatum,
stamen 1. flavum. Undique in stagnis littoris Charamis Malae nec non campi Ipios”. However this material appears to
have been lost. It was regarded by Greuter et al. (1984) as probably a synonym of C. stagnalis without explanation; but
a plant with linear leaves which is flowering under water will not be C. stagnalis, and it must be treated as unresolved
unless a specimen can be found. Callitriche stagnalis was reported by Rechinger (1943) from Lesvos, however this
material was re-determined as C. brutia var. naftolskyi (see Appendix). Therefore, the presence of C. stagnalis on
Lesvos is doubtful (see also Bazos & Yannitsaros 2004). Callitriche stagnalis is very rare in the Aegean, being reported
from the islands of Skiathos (Economidou 1969), Kos (Hansen 1980) and Ikaria (Christodoulakis 1996), but none of
these records has been critically confirmed. Schotsman & Mathez (1983) and Greuter et al. (1984) consider that it
probably does not occur in the East Aegean Islands and it is best to treat all reports from the region as erroneous unless
they can be confirmed. A plant collected by Edmondson (E) is C. truncata Guss. subsp. truncata, which settles the
presence of this taxon on the island. Thus the only Callitriche taxa which can be confirmed as occurring on Lesvos
are C. brutia var. brutia, C. brutia var. naftolskyi, C. lusitanica, C. truncata subsp. occidentalis (Rouy 1910: 186)
Schotsman (1986: 153) and C. truncata subsp. truncata.
Callitriche lusitanica was found in Sardinia when sites from which C. truncata subsp. truncata had erroneously
been reported were visited, while C. brutia var. naftolskyi was found in Sicily when visiting sites with records of the
same taxon or of C. brutia var. brutia (Pasta et al. 2008). This suggests that misidentification of these taxa is common.
Review of material at BM, CGE, E, G. K, SS and STU has located C. truncata subsp. truncata material from a single
site in Sardinia, the region around Palermo in Sicily, Barbalias (a small island off the NE coast of Lesvos), and a number
of locations in Syria. The sites around Palermo are very likely to have been destroyed due to coastal development since
these collections were made. A second site in Sicily from which this taxon has been recorded (Minissale & Sciandrello

98 • Phytotaxa 313 (1) © 2017 Magnolia Press LANSDOWN ET AL.

2016) was not visited but it would be valuable to confirm the record. No Callitriche was found at the site on Barbalias
when it was visited by IB in the 1990s. It is not currently possible to assess the potential for survival of populations in
Syria. It is clear that C. truncata subsp. truncata is very rare, however the distinction of subsp. truncata from subsp.
occidentalis is difficult from herbarium specimens and there is a need for fieldwork to locate extant populations.
Records of C. regis-jubae from Italy are based on a specimen collected from near “R.s. Giovanni, 18 km au NW
de Olbia, 30 m a.s.l., 21 April 1971, Ast & Boischler” (P), determined by H.D. Schotsman, and on material from the
island of Capraia (Lansdown et al. 2014). The latter material has been redetermined as C. brutia var. naftolskyi and it
seems wise to consider the occurrence of C. regis-jubae unconfirmed in Italy until the specimen at P can be reviewed.
Callitriche brutia var. brutia and var. naftolskyi, C. lenisulca Clavaud, C. lusitanica, C. obtusangula, C. stagnalis, C.
truncata subsp. occidentalis and subsp. truncata have been confirmed from Sardinia, but it is likely that records of
C. brutia var. hamulata (Kützing ex W.D.J. Koch 1835: 246) Lansdown (2006b: 113), C. palustris L., C. platycarpa
Kützing (1832: 174) and C. regis-jubae (Pignatti 1982, Schotsman & Mathez 1983, Arrigoni 2013) are erroneous.
Herbarium material from Sicily has not been thoroughly reviewed, but C. brutia var. naftolskyi, C. lusitanica, C.
obtusangula and C. truncata subsp. truncata have been confirmed from the island.
As noted by Schotsman (1967) and Schotsman & Mathez (1983), C. brutia var. hamulata probably does not
occur in the Mediterranean basin. RVL has seen one specimen, collected by Attila Mesterházy from a small pool in the
Gorgo del Drago, Godrano, Sicily which appeared to be this variety. However, given the potential significance of such
a record, we cannot consider it as confirmed, and therefore records of C. brutia var. hamulata from the Mediterranean
region should be treated as doubtful unless confirmed through chromosome counts or molecular analysis.
The discovery that C. lusitanica has a wider range than previously thought clearly shows that the genus is
inadequately understood in the region. Combined with the recent discovery of C. pulchra Schotsman (1967: 40) in
Cyprus (Lansdown et al. 2016), together with clarification of the taxonomic position and distribution of C. brutia var.
naftolskyi, it serves to highlight the need for further critical review of herbarium material, as well as for field surveys
and dissemination of identification guidance.

Key to Callitriche species recorded from the Mediterranean region

(Note that non-flowering plants cannot be identified reliably)

1. Pollen yellow; anthers on filaments >1 mm long . ............................................................................................................................ 2

– Pollen colourless; anthers subsessile or on filaments ≤ 1 mm long ................................................................................................ 14

2. Fruit clearly winged at least at the apex ............................................................................................................................................ 3

– Fruit unwinged ................................................................................................................................................................................ 11

3. Fruit winged only at the apex; mature fruit black .............................................................................................. Callitriche palustris
– Fruit winged throughout; mature fruit black, brown or greyish . ...................................................................................................... 4

4. Bracts present .................................................................................................................................................................................... 5

– Bracts lacking .................................................................................................................................................................................... 9

5. Some fruit pedunculate; mature fruit maroon or dark brown . .......................................................................................................... 6

– All fruit sessile; mature fruit pale brownish or greyish . ................................................................................................................... 7

6. Fruit as high as or higher than wide, < 1.2 mm wide; style > 2 mm long; peduncle < 2 mm long . .................. Callitriche mathezii
– Fruit wider than high, ≥ 1.2 mm wide; style < 1 mm long; peduncle ≤ 55 mm long . ................................... Callitriche regis-jubae

7. Testa cells punctate; some bracts forked.............................................................................................................. Callitriche cribrosa

– Testa cells cross shaped; bracts simple . ............................................................................................................................................ 8

8. Pollen bluntly triangular; mature fruit pale brownish ..................................................................................... Callitriche platycarpa
– Pollen ellipsoid; mature fruit greyish ................................................................................................................ Callitriche stagnalis

9. Flowers solitary; fruit < 1 mm wide and long .................................................................................................. Callitriche lusitanica
– Male and female flowers in most axils; fruit ≤ 1 mm wide and long . ............................................................................................ 10

10. Fruit sessile . ...................................................................................................................................................... Callitriche terrestris

– Fruit pedunculate .................................................................................................................................................. Callitriche deflexa

11. Fruit pedunculate ............................................................................................................................................. Callitriche mouterdei

– Fruit sessile....................................................................................................................................................................................... 12

Callitriche (Plantaginaceae) Phytotaxa 313 (1) © 2017 Magnolia Press • 99

12. Fruit higher than wide; pollen elongate . ...................................................................................................... Callitriche obtusangula
– Fruit as wide as high; pollen ellipsoid . ........................................................................................................................................... 13

13. Different stem branches typically with either male or female flowers, with few if any of the other sex ..... Callitriche cophocarpa
– Pairs of male and female flowers alternate along shoots .................................................................................. Callitriche lenisulca

14. Fruit with mericarps divergent; cross shaped when viewed from above; stem and leaf scales lacking ......................................... 15
– Fruit with mericarps parallel; stem and leaf scales present . ........................................................................................................... 17

15. Fruit unwinged ................................................................................................................... Callitriche truncata subsp. occidentalis

– Fruit winged .................................................................................................................................................................................... 16

16. Wing broad; peduncle recurved . ......................................................................................................................... Callitriche pulchra

– Wing narrow; peduncle various but not recurved .................................................................... Callitriche truncata subsp. truncata

17. Fruit wider than high ...................................................................................................................... Callitriche brutia var. naftolskyi
– Fruit as high as or higher than wide..................................................................................................................................................18

18. Fruits sessile to shortly pedunculate (< 2 mm); leaf apices shallowly to deeply notched, with notch broad, irregular or regular* ....
......................................................................................................................................................... Callitriche brutia var. hamulata
– Fruits subsessile to long-pedunculate (usually > 2 mm); leaf apices shallowly to deeply notched, with notch narrow and often
irregular* ............................................................................................................................................... Callitriche brutia var. brutia

* Intermediates cannot reliably be separated.

Acknowledgements

Fieldwork by RVL in Sardinia and Sicily was made possible by grants from the Mohammed bin Zayed Species
Conservation Fund and the Botanical Research Fund of the British Museum (Natural History). We would like to thank
Ralph Becker for his Callitriche records, Attila Mesterhazy for sending specimens and Carol Drinkwater of RBGE for
providing images of Edmondson’s specimen of C. truncata subsp. truncata from Lesvos. RVL is grateful to Simonetta
Bagella and Maria Carmela Caria for allowing him to use the facilities of Sassari University to study C. brutia var.
naftolskyi, and to the curators of ATHU, BM, BR, C, CGE, FI, G, K, L, P, RAB, U and WAG herbaria for help with
access to specimens. We are also extremely grateful to Professor Manuel B. Crespo and an anonymous reviewer for
constructive comments on earlier drafts.

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APPENDIX. Material studied

Callitriche brutia var. brutia:—GREECE. Lesvos: SE of Agra, near Apothika, in small pool, 28 November 1993,
I. Bazos 1454 (ATHU!); ca. 2 km NE of Achladeri, sandy bay and saltmarsh, 3 April 1994, Nielsen & Skovgaard
9590 (C); approximately 4.5 km N of Kalloni, small temporary pond in a Pinus brutia forest, 24 April 1994, I. Bazos
1654, A. Yannitsaros 8575 (ATHU); 2–3 km ENE of Skalochori, 3 April 1994, Nielsen & Skovgaard 9615 (C); in the
locality called Mikri Limni, small shallow lake, 25 May 1994, I. Bazos 1761 (ATHU); ca. 7 km NE of crossroads to
Palios (ENE of Mandamados), in seasonal pools, 2 June 1994, I. Bazos 1907 (ATHU); Approximately ca. 2 km SSE
of Mesotopos, Dergatsidi, in semi-natural pool, 39°07’ N, 26°01’ E, 20 m a.s.l., 23 August 1996, A. Yannitsaros 9193
(ATHU!). Milos: Nomos Kikladon, Eparchia Milou, small stream near Achivadolimni, 36° 41’ 15” N, 24° 26’ 22” E,
10 m a.s.l., 21 March 2016, I. Bazos & A. Zikos 4690 (ATHU!).

Callitriche brutia var. naftolskyi:—GREECE. Lesvos: In fossis stagni exsiccati “Megali Limni”, Insula Mytilini
(Lesbos), 18–24 May 1934, K.H. & F. Rechinger 5723 (G!) (Rechinger 1943, Greuter & Raus 1984); ca. 3.5 km
NW of Kalloni very close to Moni Limonos, wet flat places and seasonal pools, 28 April 1994, A. Yannitsaros 8649
(ATHU!). Milos: Nomos Kikladon, Eparchia Milou, near Psathadika, small stream and temporary ponds, 36° 43’ 10”
N, 24° 21’ 35” E, 140 m a.s.l., 9 March 2016, I. Bazos & A. Zikos 4696 (ATHU!). ISRAEL. Sharon Plain, NE of Tel
Aviv, 23 April 1927, Naftolsky 01853 (HUJ). ITALY. Capraia: pozzetta sul lato est del sentierio da S. Rocco verso
S. Stefano, Isola di Capraia (Livorno), Toscana, UTM: 33T 568.4765, 125 m, 25 April 2013, A. Soldano 16260 (FI!)
(Lansdown et al. 2014). Sardinia: Marais sales pres Villacidro, March 1854, Huet de Pavilion (BM!, P). Dominant in
ditch, east of Perdasdefogu, 39º 40’ 14.8” N, 09º 30’ 22.5” E, 23 April 2016, R.V. Lansdown; Poached hollow along
track, Paùli Piccia (Tuili, VS), 39º 44’ 18.5” N, 08º 57’ 27.3” E, 23 April 2016, R.V. Lansdown; Open, very stony and
dry pool, Paùli Maiori (Tuili, VS), 39º 44’ 35.2” N, 08º 57’ 31.2” E, 23 April 2016, R.V. Lansdown; very extensive
rutted area along track, Paùli Maiori (Tuili, VS), 39º 44’ 32.2” N, 08º 57’ 31.2” E, 23 April 2016, R.V. Lansdown;
shallow hollow in rocky plateau, Monte Minerva (Villanova Monteleone, SS), 40º 26’ 28.1” N, 08º 32’ 24.6” E, 24
April 2016, R.V. Lansdown; shallow hollow in rocky plateau, Monte Minerva (Villanova Monteleone, SS), 40º 26’
31.4” N, 08º 32’ 34.6” E, 24 April 2016, R.V. Lansdown; shallow hollow in rocky plateau, Monte Minerva (Villanova
Monteleone, SS), 40º 26’ 32.0” N, 08º 32’ 24.8” E, 24 April 2016, R.V. Lansdown; large shallow pool beside track,
Monte Minerva (Villanova Monteleone, SS), 40º 26’ 32.5” N, 08º 32’ 26.7” E, 24 April 2016, R.V. Lansdown. Sicily:
Palermo, 1854 (K!); seasonal pool, Buccheri, Siracusa Province, 37° 7’ 12.51” N, 14° 51’ 22.60” E, 27 April 2016,
R.V. Lansdown & A. Troia; cupular pool in karstic limestone, Castello della Pietra, Castelvetrano, Trapani Province,
37°40’1.38”N, 12°53’33.70”E, 28 April 2016, R.V. Lansdown & A. Troia. LIBYA. Drying pool between El Merg

Callitriche (Plantaginaceae) Phytotaxa 313 (1) © 2017 Magnolia Press • 103

(Barce) & Holmetta, Benghazi Province, 14 April 1939, N.D. Simpson 39607 (BM!); Lamluda, Derma Province, 11
April 1939, N.J. Sandwith 2571, N.D. Simpson 39512 (BM!). MOROCCO. Daya au sud de Marchaud, 45 m, 28 April
1927, E. Jahandiez 35 (BM!, G!); Daya a l’W de la route Chemmaiya–Chichaoua au N. de Bled Haroussia, 6 March
1956, Nègre 4876 “rel. à 2368” (RAB). SYRIA. 1 km N of Quneitra, 7 May 1953, P. Mouterde P 213 (G!).

Callitriche lusitanica:—GREECE. Lesvos: between Parakila and Agra, crossroads to Apothica, 39° 07’ 31”N, 26°
05’ 11”E, ca. 130 m, 25 April 1994, Bazos 1683 (ATHU); approximately 3.5 km NE of Sigri, temporary pond, 39° 13’
27” N, 26° 52’ 43” E, 16 April 2015, I. Bazos & A. Zikos 4670 (ATHU!). ISRAEL. Vernal pool, 10 km NNE of Ramat
Magshimim, Golan, 18 June 1984, A. Danin s.n. (BR! 24932653). ITALY. Sardinia: Ollastu River, 39º 20’ 52.5” N,
09º 27’ 08.4” E, 23 April 2016, R.V. Lansdown; backwater of Rio Picocca, 39º 21’ 18.0” N, 09º 29’ 34.7” E, 23 April
2016, R.V. Lansdown; broad shallow channel flowing through pasture, Fiume Temo, 40º 23’ 54.7” N, 08º 28’ 34.4” E,
24 April 2016, R.V. Lansdown; mainly terrestrialised on rocks on bedrock-dominated river, 40º 31’ 31.4” N, 08º 28’
34.4” E, 24 April 2016, R.V. Lansdown. Sicily: robably permanent pool in the Bosco Pisano area, Buccheri, Siracusa
Province, 37° 10’ 32.67” N, 14° 51’ 58.33” E, 3 April 2010, A. Troia & F. Marrone.

Callitriche terrestris:—SPAIN. Andalucía, Córdoba, between stones in street, 119 m a.s.l., 37° 52.76’ N, 04° 46.60’
W, 4 May 2016, J.J. & E.H. Wieringa 8676 (WAG!, BM!).

Callitriche truncata subsp. truncata:—ALGERIA. Prov. de Constantine, July 1861, E. Cosson 3 (P!). GREECE.
Barbalias: In a tank of fresh water by the shepherds’ cabin, Nomos Lesvou, Eparchia Mitilinis, 39° 19’ N, 26° 26’ E,
6 May 1978, Edmondson & McClintock 2468 (E) (Edmondson 1982). ITALY. Sardinia: In rivulis prope Pulam, April
[no year given] V.I. Müller (CGE!, K!, L!). Sicily. Palermo, s.d., Todaro (CGE!); Palermo a Mondello, s.d., Todaro
(P!); in calcarcis hyeme inundatis, Palermo, 21 April 1882, Todaro 623 (K!, P!, U!); in water, Palermo, s.n., 1853 (K!);
in locis hieme inundatis Vergine Maria prope Panormum, 15 March 1856, E. & A. Huet du Pavillon (K!, P!, WAG!);
April 1903, A. Prior (K!).

104 • Phytotaxa 313 (1) © 2017 Magnolia Press LANSDOWN ET AL.

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