Webbia: Journal of Plant Taxonomy and Geography, 2018

VOL. 73, NO. 1, 55–61

https://doi.org/10.1080/00837792.2018.1443552

Notes on the genus Callitriche (Plantaginaceae, Callitricheae) in South America,

and an identification key for Brazil
Gustavo Hassemera  and Richard V. Lansdownb,c 
a
Statens Naturhistoriske Museum, Københavns Universitet, Copenhagen, Denmark; bArdeola Environmental Services, Stroud, Gloucestershire,
UK; cRoyal Botanic Gardens, Kew, UK

ABSTRACT ARTICLE HISTORY

Callitriche is a genus of small herbs, ranging from aquatic to amphibious and terrestrial, Received 10 December 2017
the taxonomy of the genus is complicated and still unresolved, and knowledge of species Accepted 19 February 2018
distribution is still deficient. The present work aims to contribute to the taxonomic and
KEYWORDS
geographical knowledge of Callitriche in South America by presenting the first records of C. Callitriche lechleri; Callitriche
terrestris subsp. subsessilis in the Brazilian states of Rio de Janeiro, Santa Catarina and Sergipe, stagnalis; Callitriche terrestris
and by presenting evidence that C. lechleri and C. stagnalis, which have been reported for Brazil subsp. subsessilis; distribution
in taxonomic databases, do not occur in this country. We also lectotypify C. lechleri, and present extension; Lamiales; new
the first records of C. stagnalis in Chile and the Falkland Islands. Finally, we provide a key to the records; typification
identification of the four species of Callitriche that occur in Brazil.

Introduction published by Funez et al. (2017), and this species was

Callitriche L. (Lamiales Bromhead, Plantaginaceae Juss.,
Callitricheae Dumort.) is a cosmopolitan genus includ-
ing c. 75 species concentrated in temperate regions
(Philbrick and Jansen 1991; Philbrick and Les 2000;
Miller 2001; Erbar and Leins 2004; Villani et al. 2004;
Bean 2007; García 2010). This genus, along with Hippuris
L., Littorella P.J.Bergius and Plantago L. is notable among
the Plantaginaceae for the much simplified morphology,
particularly of the floral parts (Hegelmaier 1864; Miller
2001; Lansdown 2006b, 2009). Pollination in Callitriche
has been shown to be particularly variable, with anemo-
phily, epihydrophily and hypohydrophily having been
reported (Philbrick and Anderson 1992; Osborn and
Philbrick 1994; Cooper et al. 2000; Philbrick and Les
2000).
The taxonomy of Callitriche is still very problematic,
and a great percentage of herbarium materials are either
only identified to genus or misidentified (Miller 2001;
Lansdown 2006b, 2009). The phylogenetic relationships
within the genus also remain to be clarified (Philbrick
and Jansen 1991; Philbrick and Les 2000; Prančl et al.
2014). Some regional works provide important infor-
mation on the taxonomy and distribution of this genus
in South America (Hegelmaier 1875; Dawson 1965;
Bacigalupo 1979a, 1979b; Mereles and Degen 1993;
Novara 1993; Bittrich 2002; Zuloaga et al. 2008; Souza
2010). In Brazil, two species were recorded by Souza
(2010); this number was later increased to six species in
Souza and Hassemer (2015). New distribution records
for C. deflexa A.Braun ex Hegelm. in Brazil were recently
lectotypified by Lansdown et al. (2017).
The present work aims to contribute to the taxonomic
and geographical knowledge of Callitriche in South
America by presenting the first records of C. terrestris
subsp. subsessilis (Fassett) Bacigalupo in the Brazilian
states of Rio de Janeiro, Santa Catarina and Sergipe, and
by presenting evidence that C. lechleri (Hegelm.) Fassett
and C. stagnalis Scop., which have been reported for
Brazil in taxonomic databases, do not occur in this coun-
try. We also lectotypify C. lechleri, and present the first
records of C. stagnalis in Chile and the Falkland Islands.
Finally, we provide a key to the identification of the four
species of Callitriche that occur in Brazil.
Material and methods
We revised the Callitriche collections of ASE, BM, C,
EAC, EFC, F, FI, FLOR, FT, FURB, G, GB, GH, HBR,
HRB, HURB, K, MA, MBM, MO, NMW, NY, P, PA,
STU, TEPB, TUB, UESC and US (acronyms according
to Thiers 2017+). The conservation status assessments
follows the International Union for Conservation of
Nature (2012, 2017) criteria.
Taxonomic treatment
1. New records of Callitriche terrestris subsp. subsessilis
Callitriche terrestris subsp. subsessilis (Fassett)
Bacigalupo, Flora Ilustrada de Entre Ríos 5: 577–578.
1979. Figure 1.

CONTACT  Gustavo Hassemer  gustavonaha@gmail.com

© 2018 Dipartimento di Biologia, Università di Firenze

Published online 29 Mar 2018

56   G. HASSEMER AND R. V. LANSDOWN
Figure 1.  Herbarium specimen of Callitriche terrestris subsp.
subsessilis (G. Viana 626, ASE-2736) collected in Riachão
do Dantas, Sergipe state, north-eastern Brazil. Copyright:
Universidade Federal de Sergipe.
Type: Colombia, Norte de Santander, east of Bata, val-
ley of Rio Chitagá, south-west of Pamplona, alt. 6500
ft, 13 October 1944, N.C. Fassett 25932 (holotype,
MO-1607122!; isotypes, GH-00048930!, NY-00248743!,
US-1989482, barcode 00095624!, US-2166226, barcode
00609860!).
(≡) C. deflexa var. subsessilis Fassett, Rhodora 53: 150.
1951.
Description
Small terrestrial herbs. Sparsely distributed, rather
inconspicuous peltate scales present on stems and leaves.
Leaves isomorphic, obovate to spathulate, slightly fused
at their bases, 2.0–4.0 × 0.5–2.0 mm, entire, apex obtuse.
Bracts absent. One staminate and one pistillate flower
borne in the same leaf axil, or more rarely flowers iso-
late. Staminate flowers with a pedicel, c. 0.5 mm long;
anthers 0.10–0.15 mm long, yellow, more or less reni-
form. Pistillate flowers and fruits with an inconspicu-
ous pedicel, growing to 0.1–0.4 mm long. Mature fruits
wider than long, 0.5–0.8 × 0.7–1.0 mm (including wing),
compressed, not thickened or flared at the base, brown,
surface reticulate, faces plane; wing narrow (<10% width
of mericarp), conspicuous with magnification, evenly
surrounding the mericarps.
Illustrations
Figure 294(A–E) in Bacigalupo (1979a); figure 7(A–G)
in Bacigalupo (1979b); figure 2 in Mereles and Degen
(1993).
Distribution
Callitriche terrestris subsp. subsessilis is the terres-
trial Callitriche taxon most widely distributed in the
Americas: it occurs from central Mexico to central
Argentina (Fassett 1951; Bacigalupo 1979a, 1979b;
Bittrich 2002). In Brazil, this taxon had been hitherto
recorded in Bahia, Minas Gerais, Pernambuco and São
Paulo states (Fassett 1951; Bittrich 2002). Here we pres-
ent the first records of this taxon in Rio de Janeiro, Santa
Catarina and Sergipe states (Figure 2).
Habitat
Grows on humid soil, especially close to water bod-
ies, in shaded spots and in areas prone to occasional
inundation.
Conservation status
Least concern (LC). This subspecies has a broad distri-
bution, from southern North America to central South
America, and does not appear to be threatened.
Notes
This is the only subspecies of C. terrestris Raf. known to
occur in Brazil.
Additional material examined
S.l., s.d., A.F.M. Glaziou 8077 (C). Argentina, Corrientes,
Empedrado, Estancia La Yela, 7 October 1980, T.M.
Pedersen 12938 (C); Mburucuyá, Estancia Santa
Teresa, 25 November 1959, T.M. Pedersen 5298 (C);
Formosa, August 1919, Gonzalez 3240 (GH-01147956
[paratype]); Jujuy, Zapla a San Salvador de Jujuy, 9
November 1974, A. Burkart et al. 30556 (P-04214914);
Salta, San Martín, Laguna La Yuntada, 20 km al NW de
Tartagal, c. 700 m, 13 September 1983, L.J. Novara 3705
(MO-3787435); Tucumán, Leales, 300 m, October 1919,
S. Venturi 482 (GH-01147935 [specimen B]); Capital,
Parque 9 de Julio, 450 m, August 1923, S. Venturi 2052
(MA-788035). Bolivia, s.d., G. Mandon 1456 (GH-
01147936, MO-1913778); Tarija, July–August 1846,
M.H.A. Weddell 4038 (P-04214848). Brazil, s.d., A.
Regnell III-1762 (P-04214852); 1844, M.H.A. Weddell
1007 (P-04214893); locis humidis hortensibus, 900 m,
2 October 1914, P.K.H. Dusén s.n. (GH); Bahia, s.d.,
P. Salzmann (MO-1913775 [paratype]); Minas Gerais,
Caldas, 22 September 1867, A. Regnell III-1578

Figure 2. Recorded distribution of Callitriche terrestris subsp. subsessilis in Brazil. Green dots and areas shaded in green (in the case
of lack of more precise geographical information) indicate the previously-known distribution, whereas red diamonds indicate the
new records.
(F [isoparatype], P-04214854 [isoparatype], P-04214858
[isoparatype]); Ouro Preto, 1894, M. Gomes 26804 (F);
Pernambuco, moist sand, among rocks by nearly dry
streamlet through dry scrub (caatinga), Bello Jardim,
Serra do Genipapo, 625 m, 14–15 November 1924, A.
Chase 7710 (US); Rio de Janeiro, 1816–1821, A.F.C.P.
de Saint-Hilaire 486 (P-04214916); Petrópolis, Sto.
Antonio, 24 September 1875, A.F.M. Glaziou 8077
(P-04214894); Rio de Janeiro, weed in garden, Quinta
de Boa Vista, 11 July 1967, F.R. Fosberg 48526 (US);
Santa Catarina, s.d., J.F.T. Müller 413 (K); Tubarão,
no terreno humido de Conconhaz perto de Tubarão,
September 1889, E. Ule 1257 (P-04214851, STU);
Sergipe, Riachão do Dantas, mata de Riachão, em
caatinga hipoxerófila, 27 August 1982, G. Viana 626
(ASE-2736; see Figure 1). Colombia, Cundinamarca,
Bogotá, Campos de la Universidad Nacional de
Colombia, 2600  m, 10 April 1986, J.L. Fernández
A. 7798 (MA-508417); Santander, Vélez, in patio of
Hotel Santander, 7120 ft, 17 April 1944, N.C. Fassett
25071 (MO-1607112 [paratype], US-2166143, barcode
00609859 [isoparatype]); Valle del Cauca, Bosque
de San Antonio, west of Cali, near television tower,
1950–2050  m, 15 July 1984, A. Gentry et al. 48173
WEBBIA: JOURNAL OF PLANT TAXONOMY AND GEOGRAPHY   57
(MO-3189994). Ecuador, lièux humìdes des forêts,
entre Guayanda et Bodegas, November 1856, Remy s.n.
(P-04214867, P-04214869); Cotopaxi, Tenefuerste, Río
Pilalo, km 52–53, Quevado, Latacunga, 750–1300 m,
21 February 1982, C.H. Dodson and A.H. Gentry 12767
(MO-2944171). Mexico, 1841–1843, F.E. Liebmann
1088 (C); México, Temascaltepec, Nanchititla, cliff,
10 November 1933, G.B. Hinton 4988 (GH-01147937
[paratype], P-04214895 [isoparatype]); Veracruz,
Jalapa, s.d., C.G. Pringle 7817 (GH-01147934).
Paraguay, s.d., P. Jörgensen 4509 (MO-971747 [para-
type]); s.d., C. Osten s.n. (MO-5599203); June 1915,
T. Rojas 1255 (MO-5599204); Luque, sur la terre
humìde, 7 June 1874, B. Balansa 2352a (P-04214847);
L’assomption, sur le bord des mares, September 1875, B.
Balansa 2352 (P-04214846, P-05109233); L’assomption,
sur le bord des mares, December 1876, B. Balansa 2353
(P-04214845). Peru, Cajamarca, Contumazá, Yetón
(San Benito-Guzmango), 1800  m, 12 June 1983, A.
Sagástegui 10554 (MO-5171459, MO-5182435).
2. Notes on Callitriche stagnalis in South America
Callitriche stagnalis Scop., Fl. Carniol., 2nd ed. 2: 251.
1772
58   G. HASSEMER AND R. V. LANSDOWN
Type: UK, Ceredigion, Aberleri Fields, Borth, UK
Ordnance Survey grid 22/61209160, 20 July 1998, A.O.
Chater s.n. (neotype NMW!; designated by Lansdown
2006a, p. 108).
Illustrations
Figures on p. 81 and 83 in Lansdown (2008).
Distribution
Callitriche stagnalis is native to Europe, from the Azores
to Iceland and east to the western side of the Ural
Mountains in Russia (Lansdown 2008). It also occurs
as an alien (naturalised) in North America (Philbrick
et al. 1998), Australia (Orchard 1980), New Caledonia
and New Zealand (Mason 1959).
Conservation status
Least concern (LC). This species has a broad distribution
and does not appear to be threatened.
Notes
Callitriche stagnalis has been reported to occur in Brazil
(Kaehler 2014; Souza and Hassemer 2015) based on one
collection from Curitiba, Paraná state, southern Brazil
Figure 3. Herbarium specimen of Callitriche rimosa (J. Cordeiro
and E. Barboza 794, C) collected in Curitiba, Paraná state,
southern Brazil. Copyright: Københavns Universitet.
(J. Cordeiro and E. Barboza 794, C!, MBM!; see Figure
3). However, careful examination of these specimens has
revealed that they belong instead to C. rimosa Fassett.
Since there is no other record of C. stagnalis in Brazil,
and this species is not recorded in neighbouring coun-
tries, we believe that it does not occur in this country,
and therefore we propose its removal from the flora of
Brazil.
With our revisions of herbarium collections we dis-
covered the first records of C. stagnalis in Chile (T.G.
Lammers et al. 7503, F-2068478!; see Figure 4) and the
Falkland Islands (R. Lewis s.n., K-000299348!). In both
cases there is clear indication that the species occurs in
anthropic environments, i.e. areas under strong influ-
ence of human activities. These are the first verified
records of C. stagnalis in South America; based on what
happened in North America and Australasia, we believe
that this species will probably expand its distribution in
temperate areas of South America.
New records
Chile, Biobío, Concepción province, Concepción, Estero
Lenga, 2.5 km west of Avda. Gran Bretaña on Avda. Los
Golondrinas, just beyond the Petrox refinery, submersed
Figure 4.  Herbarium specimen of Callitriche stagnalis (T.G.
Lammers et al. 7503, F-2068478) collected in Concepción
province, Chile. Copyright: Field Museum of Natural History.

in shallow pools in grazed salt marsh, 22 October 1990,
T.G. Lammers et al. 7503 (F-2068478; see Figure 4). UK,
Falkland Islands, East Falkland, Stanley, ditches/drain-
age, 2 March 2009, R. Lewis s.n. (K-000299348).
3. Lectotypification of Callitriche lechleri and its removal
from the flora of Brazil
Callitriche lechleri (Hegelm.) Fassett, Rhodora 53: 191–
194, Tab. 1174(18). 1951
Type: Chile, O’Higgins, Rancagua, May–June 1828,
C.L.G. Bertero 142 (lectotype designated here,
GH-00048932!; isolectotypes, BM!, MO-1913783!,
NY-00248745!, P-04214934!). Figure 5.
(≡) C. marginata var. lechleri Hegelm., Verh. Bot. Vereins
Prov. Brandenburg 9: 12–13. 1867.
Illustrations
Tab. 1174(18) in Fassett (1951); figure 296 in Bacigalupo
(1979a); figure 4 in Bacigalupo (1979b).
Distribution
Argentina, Chile and Uruguay (Fassett 1951; Bacigalupo
1979a, 1979b; Zuloaga et al. 2008).
Figure 5. Lectotype of Callitriche marginata var. lechleri (C.L.G.
Bertero 142, GH-00048932). Copyright: Harvard University.
WEBBIA: JOURNAL OF PLANT TAXONOMY AND GEOGRAPHY   59
Conservation status
Least concern (LC). This species has a relatively broad
distribution in southern South America and does not
appear to be threatened.
Notes
The protologue of C. marginata var. lechleri (Hegelmaier
1867, p. 12–13) provided the following information
on the type: ‘Chile (Santiago, Philippi, K. K. W. H.
[Kaiserlich-königliches Wiener Herbarium (= W herbar-
ium)]; Valdivia, Lechl. pl. chil. N. 260; Quillota, Bertero,
Hb. Rich.; Runcagua, Bert. N. 142, Herb. Steud.) und
Peru (Ayapata, Lechl. pl. Peruv.)’. Later, Fassett (1951,
p. 192) indicated the gathering C.L.G. Bertero 142 as
the type for this name, listing specimens kept at three
herbaria as co-types: ‘Chile: Rancagua, May, June, 1828,
Bertero 142 (MO, GH, NY – Cotypes)’. Because of the
importance of Fassett’s work on Callitriche, and aiming at
conserving the current application of Callitriche names
as much as possible, we decided to select a lectotype for
C. marginata var. lechleri among specimens kept at these
three herbaria cited by Fassett (1951). Following this cri-
terion, we were able to locate the following original spec-
imens: GH-00048932, MO-1913783 and NY-00248745.
We designate here the specimen GH-00048932 (Figure
5), which is the best among these three syntypes, as the
lectotype of C. marginata var. lechleri.
Callitriche lechleri has been reported to occur in
Brazil (Souza and Hassemer 2015) based on one collec-
tion from Bom Retiro, Santa Catarina state, southern
Brazil (L.B. Smith and R.M. Klein 7854, P-04214910!; see
Figure 6). This specimen was originally identified as C.
oblongicarpa Fassett (a synonym of C. lechleri, according
to Bacigalupo (1979a, 1979b) and Zuloaga et al. (2008))
by Lyman Bradford Smith. However, careful examina-
tion of this specimen has revealed that it belongs instead
to C. rimosa. Since this was the only record of C. lechleri
in Brazil, we believe that this species does not occur in
this country, and therefore we propose its removal from
the flora of Brazil.
Key to the identification of Callitriche in Brazil
Note: state-level distributions are presented inside brack-
ets. State abbreviations: BA: Bahia; MG: Minas Gerais;
PE: Pernambuco; PR: Paraná; RJ: Rio de Janeiro; RS:
Rio Grande do Sul; SC: Santa Catarina; SE: Sergipe; SP:
São Paulo.
1. Mature fruit black, up to 1 mm wide ...................... 2
Mature fruit brown, 0.8–1.5 mm wide [PR; RS; SC; SP]
................................................................ C. rimosa Fassett
2. Bracts absent. Fruit winged ........................................ 3
Bracts present at least subtending flowers but caduceus.
Fruit unwinged [RS] ..................... C. heterophylla Pursh
60   G. HASSEMER AND R. V. LANSDOWN
Figure 6. Herbarium specimen of Callitriche rimosa (L.B. Smith
and R.M. Klein 7854, P-04214910) collected in Bom Retiro, Santa
Catarina state, southern Brazil. Copyright: Muséum National
d’Histoire Naturelle.
3. Some fruit on all plants long-pedicellate; pedicel up
to 6.6 mm. Mature fruit widest at mid-point; fruit faces
slightly convex [PR; RJ; RS; SC; SP] .................................
......................................... C. deflexa A.Braun ex Hegelm
Fruit subsessile or on pedicel up to 0.5 mm. Mature fruit
widest toward apex; fruit faces plane [BA; MG; PE; RJ;
SC; SE; SP] ...........................................................................
...... C. terrestris subsp. subsessilis (Fassett) Bacigalupo
Acknowledgements
Thanks are due to Marta Cristina Vieira Farias (Universidade
Federal de Sergipe) for supporting this study; to the curators
and staff of all the herbaria cited in Material and Methods
for receiving us during our studies, for the loan of herbarium
specimens, for providing images of herbarium specimens
and for providing important information to this study; and to
anonymous reviewers for contributing to improve this paper.
Gustavo Hassemer also acknowledges three SYNTHESYS
grants for visits to K, MA and P herbaria (processes
GB-TAF-5317, ES-TAF-6610 and FR-TAF-6529). Richard
Lansdown acknowledges a grant (no. 15259235) from the
Mohammed Bin Zayed Species Conservation Fund.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
This work was supported by SYNTHESYS Project [grant
number GB-TAF-5317, ES-TAF-6610 and FR-TAF-6529].
ORCID
Gustavo Hassemer   http://orcid.org/0000-0003-4365-6934
Richard V. Lansdown   http://orcid.org/0000-0003-0984-4552
References
Bacigalupo NM. 1979a. Callitrichaceae. In: Burkart A, editor.
Flora Ilustrada de Entre Ríos. vol 5. Buenos Aires: Instituto
Nacional de Tecnología Agropecuaria; p. 576–582.
Bacigalupo NM. 1979b. El género Callitriche en la flora
argentina. Darwiniana. 22:377–396.
Bean AR. 2007. A taxonomic revision of Callitriche L.
(Callitrichaceae) in Australia. Austrobaileya. 7:545–554.
Bittrich V. 2002. Callitrichaceae. In: Wanderley MGL,
Shepherd GJ, Giulietti AM, Melhem TS, Bittrich V,
Kameyama C, editors. Flora Fanerogâmica do Estado de
São Paulo. vol 2. São Paulo: Instituto de Botânica; p. 65–66.
Cooper RL, Osborn JM, Philbrick CT. 2000. Comparative
pollen morphology and ultrastructure of the
Callitrichaceae. Amer J Bot. 87:161–175.
Dawson G. 1965. Callitrichaceae. In: Cabrera ÁL, editor.
Flora de la Provincia de Buenos Aires. vol 4. Buenos Aires:
Instituto Nacional de Tecnología Agropecuaria; p. 125–
129.
Erbar C, Leins P. 2004. Callitrichaceae. In: Kadereit JW, editor.
The families and genera of vascular plants. vol 7. Berlin
and Heidelberg: Springer; p. 50–56. doi:10.1007/978-3-
642-18617-2_5.
Fassett NC. 1951. Callitriche in the New World. Rhodora.
53:137–155, 161–182, 185–194, 209–222, plates 1167–1175.
Funez LA, Hassemer G, Ferreira JPR, Bones FLV, dos Santos
AP. 2017. Fifty-five new records of vascular plants, and
other discoveries for the flora of Santa Catarina, southern
Brazil. Webbia. 72:221–275. doi:10.1080/00837792.2017.1
369303.
García P. 2010. Callitriche L. In: Benedí C, Rico E, Güemes
J, Herrero A, editors. Flora Iberica. vol 12. Madrid: Real
Jardín Botánico; p. 497–513.
Hegelmaier CF. 1864. Monographie der Gattung Callitriche.
Stuttgart: Ebner & Seubert.
Hegelmaier CF. 1867. Zur Systematik von Callitriche.
Verhandlungen des botanischen Vereins für die Provinz
Brandenburg und die angrenzenden Länder. 9:1–41.
Hegelmaier CF. 1875. Callitrichineae. In: von Martius CFP,
Eichler AW, editors. Flora Brasiliensis. vol 13. Munich and
Leipzig: F. Fleischer; p. 1–16, plate 1. doi:10.5962/bhl.title.454.
International Union for Conservation of Nature. 2012. IUCN
red list categories and criteria. version 3.1. 2nd ed. Gland:
IUCN.
International Union for Conservation of Nature. 2017.
Guidelines for using the IUCN red list categories and
criteria. version 13. Gland: IUCN.
Kaehler M. 2014. Plantaginaceae. In: Kaehler M, Goldenberg
R, Evangelista PHL, Ribas OS, Vieira AOS, Hatschbach
GG, editors. Plantas Vasculares do Paraná. Curitiba:
Universidade Federal do Paraná; p. 161.
Lansdown RV. 2006a. Notes on the water-starworts
(Callitriche) recorded in Europe. Watsonia. 26:105–120.
Lansdown RV. 2006b. The genus Callitriche (Callitrichaceae)
in Asia. Novon. 16:354–361. doi:10.3417/1055-3177(2006)
16[354:tgccia]2.0.co;2.

Lansdown RV. 2008. Water starworts of Europe. BSBI
Handbook no. 11. London: Botanical Society of the British
Isles.
Lansdown RV. 2009. Nomenclatural notes on Callitriche
(Callitrichaceae) in North America. Novon. 19:364–369.
doi:10.3417/2008030.
Lansdown RV, Glen R, Hassemer G. 2017. Notes on the genus
Callitriche (Plantaginaceae) in South Africa. Phytotaxa.
317:219–225. doi:10.11646/phytotaxa.317.3.6.
Mason R. 1959. Callitriche in New Zealand and Australia.
Aust J Bot. 7:295–327. doi:10.1071/bt9590295.
Mereles F, Degen R. 1993. Callitrichaceae. In: Spichiger R,
Ramella L, editors. Flora del Paraguay, vol. 18. Geneva:
Conservatoire et Jardin botaniques de la Ville de Genève;
and St. Louis: Missouri Botanical Garden; p. 1–13.
Miller NG. 2001. The Callitrichaceae in the southeastern
United States. Harvard Pap Bot. 5:277–301.
Novara LJ. 1993. Flora del Valle de Lerma: Callitrichaceae
Link. Aportes Botánicos de Salta, serie Flora. 1(21):1–5.
Orchard AE. 1980. Callitriche (Callitrichaceae) in South
Australia. J Adel Bot Gard. 2:191–194.
Osborn JM, Philbrick CT. 1994. Comparative pollen structure
and pollination biology in the Callitrichaceae. Acta Bot
Gallica. 141:257–266. doi:10.1080/12538078.1994.10515157.
Philbrick CT, Anderson GJ. 1992. Pollination biology in the
Callitrichaceae. Syst Bot. 17:282–292. doi:10.2307/2419523.
Philbrick CT, Jansen RK. 1991. Phylogenetic studies of North
American Callitriche (Callitrichaceae) using chloroplast
DNA restriction fragment analysis. Syst Bot. 16:478–491.
doi:10.2307/2419338.
Philbrick CT, Les DH. 2000. Phylogenetic studies in
Callitriche: implications for interpretation of ecological,
WEBBIA: JOURNAL OF PLANT TAXONOMY AND GEOGRAPHY   61
karyological and pollination system evolution. Aquat Bot.
68:123–141.
Philbrick CT, Aakjar RA Jr, Stuckey RL. 1998. Invasion and
spread of Callitriche stagnalis (Callitrichaceae) in North
America. Rhodora. 100:25–38.
Prančl J, Kaplan Z, Trávníček P, Jarolímová V. 2014.
Genome size as a key to evolutionary complex aquatic
plants: polyploidy and hybridization in Callitriche
(Plantaginaceae). PLoS ONE. 9(9):e105997. doi:10.1371/
journal.pone.0105997.
Souza VC. 2010. Plantaginaceae. In: Forzza RC et al.,
organisers. Catálogo de Plantas e Fungos do Brasil, vol.
2. Rio de Janeiro: Andrea Jakobsson Estúdio and Jardim
Botânico do Rio de Janeiro; p. 1460–1464.
Souza VC, Hassemer G. 2015. Plantaginaceae. In: Lista de
Espécies da Flora do Brasil. Jardim Botânico do Rio de
Janeiro. [accessed 2017 Dec 10]. http://www.floradobrasil.
jbrj.gov.br/jabot/floradobrasil/fb191.
Thiers B. 2017+. Index Herbariorum: a global directory of
public herbaria and associated staff. New York Botanical
Garden’s Virtual Herbarium. [accessed 2017 Dec 10].
http://sweetgum.nybg.org/science/ih.
Villani M, Marcucci R, Bassan E, Magrin M, Tornadore N.
2004. The genus Callitriche in northeastern Italy: first
systematic contribution. Israel J Plant Sci. 52:331–340.
Zuloaga FO, Morrone O, Belgrano MJ, Marticorena C,
Marchesi E. 2008. Callitrichaceae. In: Zuloaga FO,
Morrone O, Belgrano MJ, Marticorena C, Marchesi E,
editors. Catálogo de las Plantas Vasculares del Cono
Sur, vol. 2. Monographs in Systematic Botany from the
Missouri Botanical Garden, vol. 107. St. Louis: Missouri
Botanical Garden; p. 1843–1844.