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Cucumis (Cucurbitaceae) in Australia and Eastern Malesia, Including Newly Recognized Species and the Sister Species to C. melo


Abstract and Figures

Molecular phylogenetic analyses based on numerous plant accessions have shown that Cucumis comprises 25 species in Asia, the Malesian region and Australia, rather than just two as traditionally thought. Among the 25 species several are new, and here we describe four from tropical Australia. The new species C. costatus, C. queenslandicus, C. umbellatus and C. variabilis are illustrated, their distributions mapped based on 7-50 collections per species and information is provided on habitats and conservation status. We also validate C. althaeoides comb. nov. and C. argenteus comb. nov., update the description of C. picrocarpus, and provide a key to the 11 native and naturalized species of Cucumis occurring in Australia and eastern Malesia.The Australian species diversity of Cucumis, a genus that until recently was held to be essentially African, is of interest also because Australia harhours the sister species to the commercially important melon, C. melo.
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Cucumis (Cucurbitaceae) in Australia and Eastern Malesia, Including Newly
Recognized Species and the Sister Species to C. melo
Author(s): Ian R. H. Telford, Patrizia Sebastian, Jeremy J. Bruhl, and Susanne S. Renner
Source: Systematic Botany, 36(2):376-389. 2011.
Published By: The American Society of Plant Taxonomists
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Systematic Botany (2011), 36(2): pp. 376–389
© Copyright 2011 by the American Society of Plant Taxonomists
DOI 10.1600/036364411X569561
Phylogenetic reconstructions for the Cucurbitaceae have
revealed unexpected relationships for Australian species and
genera ( Kocyan et al. 2007 ). Of particular interest is the dis-
covery that Cucumis as traditionally conceived ( Kirkbride
1993 ) is paraphyletic and that it comprises 25, rather than
two, Asian, Malesian, and Australian species ( Renner et al.
2007 ; Renner and Schaefer 2008 ; Sebastian et al. 2010 ).
A particularly poorly understood African/Asian/Australian
genus found embedded in Cucumis is Mukia Arn. In the last
comprehensive treatment of the Australian Cucurbitaceae
( Telford 1982 ), Mukia was presented as containing six spe-
cies in Australia: the putatively widespread and polymorphic
M. maderaspatana (L.) M. Roem. (now Cucumis maderaspata-
nus L.), M. micrantha (F. Muell.) F. Muell. (now Austrobryonia
micrantha (F. Muell.) I. Telford), and five species not val-
idly published but assigned code names. Based on molec-
ular data, three of these unnamed species, together with
M. micrantha , form a distinct clade, described as Austrobryonia
H. Schaef., a genus of the tribe Bryonieae, an early branch
in the Cucurbitaceae ( Schaefer et al. 2008 ). Several other
new combinations required by the molecular findings, but
not concerning Australian species, have already been made
( Ghebretinsae et al. 2007a ; Schaefer 2007 ).
Molecular data obtained since 2007 for an increasingly
comprehensive sampling of Asian and Australian specimens
(over 100 accessions, mostly from herbarium collections)
have revealed that the Cucumis species treated as Mukia sp.
A and M. sp. B in the Flora of Australia ( Telford 1982 ), and
a third more recently discovered species, are close rela-
tives, in line with their morphological similarity. These three
species are sister to C. javanicus (Miq.) Ghebret. & Thulin
( Fig. 1 ). Molecular data generated for the present study also
show that Melothria argentea is a genetically distinct species
of Cucumis ( Fig. 1 ). Likewise, a species originally described
as Bryonia althaeoides by Seringe (1828) and later regarded as
a species of Mukia ( Roemer, 1846 ) or Melothria ( Nakai, 1938 )
turns out to be the sister species to C. variabilis , another new
Australian species ( Fig. 1 ). Lastly, the native Australian spe-
cies C. picrocarpus , which had long been regarded as a syn-
onym of C. melo ( Kirkbride 1993 ), instead is the sister species
to melon, C. melo ( Fig. 1 ). The geographic origin of C. melo
itself and its likely domestication in the Eastern Himalayas
region are dealt with in a related paper ( Sebastian et al. 2010 ).
Because of the great interest of melon breeders in C. picro-
carpus , we here provide an updated description of this spe-
cies based on new observations in the herbarium and field.
Lastly, we provide a key to the 11 native and naturalized spe-
cies of Cucumis now known from Australia and adjacent east-
ern Malesia.
Materials and Methods
Herbarium Work, Field Observations, Morphological Data—
Specimens in BRI, CANB, DNA, MEL, NE, PERTH and QRS (now in
CNS) were examined, and we also obtained scans of types from B, E,
G-DC, and PR. Floral measurements were taken on rehydrated dried
specimens. Fieldwork by the first two authors greatly improved under-
standing of the ecological niches of the native Australian species of
Cucumis .
Molecular Phylogenetics— The DNA sequence matrix for this paper
is modified from the one used in Sebastian et al. (2010) . That study
included 113 accessions of Cucumis and its sister group, Muellerargia
Cogn., sequenced for six chloroplast regions (the trnL intron, the inter-
genic spacers trnL-F , rpl20-rps12 , and trnS-G ; and the genes rbcL and
matK ) plus the nrDNA ITS1 and ITS2, and the intervening 5.8 S gene seg-
ment. The dataset comprised 6,202 aligned positions, and for the present
study, we added sequences for two accessions from Timor ( C. althaeoides
Zippel 107 (L): rpl20 - rps12 intergenic spacer (HQ439182); Cucumis sp.
nov. collector ignotus 1305 (L): rpl20 - rps12 and trnS - trnG intergenic
spacers (HQ439180, HQ439181) to resolve the application of the name
C. althaeoides . In the phylogeny shown here ( Fig. 1 ) we excluded most
African Cucumis and several undescribed species from mainland Asia
and from Africa. The phylogeny in Sebastian et al. (2010 : Fig. 2 ) and
the one included here ( Fig. 1 ) are not identical in their taxon sampling.
Sequencing approaches and phylogenetic methods are as described in
Sebastian et al. (2010) . Maximum likelihood (ML) analyses ( Felsenstein
1973 ) and ML bootstrap searches ( Felsenstein 1985 ) were performed using
RAxML ( Stamatakis et al. 2008 ).
Voucher information and GenBank accession numbers are listed in
Table S1 of Sebastian et al. (2010) . The alignment used for the present
study has been deposited in TreeBASE (study number S10944).
Results and Discussion
Based on the combined nuclear and plastid sequence data
and geographic provenence of the samples ( Fig. 1 ), there are
Cucumis (Cucurbitaceae) in Australia and Eastern Malesia, Including
Newly Recognized Species and the Sister Species to C. melo
Ian R. H. Telford , 1,3 Patrizia Sebastian , 2 Jeremy J. Bruhl , 1 and Susanne S. Renner 2
1 Botany & N. C. W. Beadle Herbarium, School of Environmental and Rural Science,
University of New England, Armidale, Australia
2 Systematic Botany and Mycology, University of Munich, Menzinger Str. 67, 80638 Munich, Germany
3 Author for correspondence (
Communicating Editor: Mark P. Simmons
Abstract— Molecular phylogenetic analyses based on numerous plant accessions have shown that Cucumis comprises 25 species in Asia,
the Malesian region, and Australia, rather than just two as traditionally thought. Among the 25 species several are new, and here we describe
four from tropical Australia. The new species C. costatus, C. queenslandicus , C. umbellatus , and C. variabilis are illustrated, their distri-
butions are mapped based on 7–50 collections per species, and information is provided on habitats and conservation status. We also vali-
date C. althaeoides comb. nov. and C. argenteus comb. nov., update the description of C. picrocarpus , and provide a key to the 11 native and
naturalized species of Cucumis occurring in Australia and eastern Malesia. The Australasian species diversity of Cucumis , a genus that until
recently was held to be essentially African, is of interest also because Australia harbours the sister species to the commercially important
melon, C. melo .
Keywords—Cucumis picrocarpus , melon , morphological characters , phylogenetics , sequencing herbarium specimens , taxonomy.
F ig . 1. Maximum likelihood tree for 27 species of Cucumis based on combined chloroplast and nuclear sequences; numbers at nodes are bootstrap
values 75%. The tree is rooted on Muellerargia , the sister group of Cucumis . Geographic occurrence of species is color-coded (inset), and the geographic
origin of each accession follows the species name. Species discussed in the text are marked by boxes.
eight indigenous species of Cucumis in Australia, of which
one also occurs in Timor and southern New Guinea. Cucumis
maderaspatanus does not occur in Australia ( Sebastian et al.
2010 ; our Fig. 1 ), and previous citations of that species for
Australia were based on misidentifications. Resolving the
circumscription of all biological species hiding under the
name C. maderaspatanus , however, requires an analysis of
specimens from throughout the range of this putatively
Asian/African species ( De Wilde and Duyfjes 2006 ). In the
present study, we begin this task by comparing relevant
Australian specimens and names ( Telford 1982 ), exclud-
ing those already transferred to Austrobryonia H. Schaef.
( Schaefer et al. 2008 ).
Besides the eight native species of Cucumis , three African
species have become widely naturalized in Australia, viz.
C. myriocarpus Naudin, C. zeyheri Sond. in Harv. & Sond., and
C. metuliferus E. Mey. ex Naudin, and there are also sporadi-
cally naturalized populations of cultivated melon, C. melo , a
species that likely evolved in the eastern Himalayan region
( Sebastian et al. 2010 ; contra Ghebretinsae et al. 2007b ).
Because of the potential interest for Cucumis breeders, we
include a key to the 11 native and naturalized Australian spe-
cies, following the taxonomic treatment of the four new spe-
cies, two new combinations, and an emended description of
C. picrocarpus .
Species of Cucumis native to Australia are descended from
five ancestral lineages that arrived from southeast Asia sev-
eral million years ago ( Sebastian et al. 2010 for molecular
clock estimates of divergence times in Cucumis ). One of the
five entries into Australia gave rise to C. costatus, C. queen-
slandicus, and C. umbellatus , while C. picrocarpus , C. melo , and
C. argenteus represent independent arrivals to the continent
( Fig. 1 ). The genetically distinct Australian endemic C. picro-
carpus diverged from C. melo about 3 Ma ago ( Sebastian et al.
2010 ), while Australian populations of C. melo all represent
naturalized cultivated forms . The early dates of collection and
their distribution suggest Australian C. melo to be indigenous,
perhaps dispersed in preEuropean times by the aboriginal
people as it is a wild food.
The last two Australian species, C. variabilis and C. althae-
oides, represent another small ‘radiation.’ This limited diver-
sification resembles the situation in other Cucurbitaceae
colonizations of Australia ( Schaefer et al. 2008 , 2009 ). A recent
review of vertebrate, invertebrate, and plant distribution
ranges across the Australian monsoon tropics implicates the
formation of the Carpentarian Gap, separating the Kimberley
region and Arnhem Land from the Cape York Peninsula in
the interruption of gene flow in numerous species ( Bowman
et al. 2009 ), and this event may also explain the C. costatus -
C. umbellatus divergence ( Fig. 1 ), which dates to the early
Quaternary ( Sebastian et al. 2010 ).
Cucumis queenslandicus, C. costatus and C. althaeoides
inhabit savannah communities adapted to seasonal heavy
rain and fire. They are perennials with thickened rootstocks
allowing dormancy in the dry season and rapid growth in
the wet. Cucumis umbellatus is an annual species occurring
on the Kimberley − Arnhem Land sandstone plateau escarp-
ments that are less fire-prone than savannah. Cucumis vari-
abilis and C. argenteus inhabit some of the most arid parts of
Australia and exhibit convergence vegetatively, both some-
times have leaves covered with dense, white, villous hairs
(see the color photos of these species provided with their
descriptions). Cucumis picrocarpus and the possible indige-
nous variant of C. melo grow on clay flats, such as alluvium
on flood plains, but are recorded from a variety of habi-
tats. More details about the vegetation types in which the
Australian species occur are provided following the species
Taxonomic Treatment
Key to Species of CUCUMIS in Australia and Eastern Malesia
Identification of Australian Cucumis specimens requires flowers, fruits, and seeds; vegetative material could be identified by
sequencing the ITS region and then BLASTing sequences against the complete Cucumis species sample in GenBank. The key
below is applicable only to Australian material and includes both native and naturalized species, with the introduced African
species marked by an asterisk.
1. Female flowers in fascicles, sometimes coaxillary with several males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Female flowers mostly 3–7 per axil; central Western Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. variabilis
2. Female flowers mostly 1 or 2, rarely 3 or 4 per axil; northern and inland Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Seeds verrucose; mostly coastal E Malesia, northern Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. althaeoides
3. Seeds smooth; mostly inland Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. argenteus
1. Female flowers solitary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Seeds pitted or verrucose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Fruit globose or subglobose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Fruit 1–3 seeded; fruiting pedicel > 5 cm long; male inflorescence umbellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. umbellatus
6. Fruit 6–25-seeded; fruiting pedicel <1 cm long; male inflorescences fasciculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. althaeoides
5. Fruit ellipsoidal, fusiform or ovoid-fusiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Male inflorescence racemose; fruit strongly ribbed, the ribs scabrid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. costatus
7. Male inflorescence fasciculate, rarely racemose; fruit weakly ribbed, the ribs hispid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. queenslandicus
4. Seeds smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Fruit aculeate; aculei scattered or dense, soft or rigid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Fruit globose; aculei soft, scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. myriocarpus
9. Fruit ellipsoidal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Aculei dense, soft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. zeyheri
10. Aculei scattered, rigidly pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. metuliferus
8. Fruit smooth or longitudinally ribbed, variously hairy, sometimes glabrescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Leaves unlobed or shallowly 3- or 5-lobed; lobes<one third of lamina width; fruit with short antrorse hairs;
native or naturalized . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. melo
11. Leaves deeply 5-lobed; lobes > one third of lamina width; fruit sparsely pilose with spreading hairs, glabrescent . . . . . . . C. picrocarpus
Cucumis picrocarpus F. Muell., descr. emend.
Cucumis picrocarpus F. Muell., Trans. Philos. Inst. Victoria 3: 46.
1859 as ‘picrocarpa.’
Cucumis picrocarpus requires lectotypification because no
spec imens were cited in the protologue, only the distributional
statement: “In many parts of tropical Australia” ( von Mueller
1859 ). We carry out this lectotypification, together with
another typification of a Mueller name, in an accompanying
paper (I. Telford, et al. in prep.).
Cucumis trigonus auct. non Roxb.: Bentham, Fl. Austral. 3: 317.
1863; F. M. Bailey, Queensland Fl. 696. 1900.
Cucumis melo auct. non L.: I. R. H. Telford, Fl. Australia 8:
189. 1982 p.p.; J. H. Kirkbride, Biosystematic mono-
graph of the genus Cucumis (Cucurbitaceae), p. 79.
1993 p. p.
Trailing or climbing annual herb, monoecious, hispid on
most vegetative parts; stems to 1.5 m long, ribbed. Tendrils
simple, to 8 cm long. Leaves: petiole 30–123 mm long; lamina
broadly ovate in outline, 28–140 × 26–130 mm, mucronate,
deeply palmately 5-lobed, the lobes lobulate, denticulate.
Inflorescences unisexual. Male flowers in (1–)3–8 flow-
ered fascicles or racemes; peduncles to 4 mm long; pedicels
3.5–12(–35) mm long; hypanthium narrowly campanulate,
3.8–4.3 mm long, hispid with spreading hairs; calyx lobes
linear to narrowly triangular, 1.8–2 mm long, hispid; corolla
lobes elliptic or ovate, 18–22 × 10–12 mm, mucronate, sparsely
hispid outside, particularly apically, glabrous inside, yel-
low; stamens inserted towards the base of the hypanthium
tube; anthers one 1-thecous, two 2-thecous, sigmoid, ca.
2 mm long; connective appendage ca. 1 mm long, lobed,
papillose; disc depressed globose, ca. 0.6 mm diam. Female
flowers solitary; pedicels 2–25 mm long; ovary ellipsoidal,
4–6.5 mm long, 3–3.8 mm diam, pilose with spreading hairs;
hypanthium above constriction 2–3.2 mm long; perianth
similar to male; disc annular, ca. 1 mm diam, lobed; style
0.6–1 mm long; stigmatic branches 1.5–1.7 mm long. Fruits
subglobose to ellipsoidal, 28–55 mm long, 25–40 mm diam,
sparsely pilose with simple, multicellular hairs, glabrescent,
pale green with darker longitudinal markings; fruiting pedi-
cel to 56 mm long. Seeds many, elliptic, 4–5.6 × 2.2–3 mm,
± compressed with the faces slightly convex, the remnant
of the funicle often remaining attached, appearing wing-
like, smooth, buff.
Representative Specimens Examined— AUSTRALIA. Western Australia:
3.5 km WSW of Barowana Hill, Hamersley Ranges, Trudgen 18824 (PERTH);
E side of old crossing at Fitzroy Crossing, Mitchell 3070 (CANB, PERTH);
Block 68, Sugar cane experimental area, Dept of Agriculture, Kununurra,
Aplin 6279 (K, PERTH); 100 m N of Lissadell Homestead main airstrip,
5 km NE of homestead, Mitchell 3076 (CANB, PERTH). Northern Territory:
Victoria Highway, ca. 19 km W from Victoria River crossing, Purdie 3261
(CANB, NA); Sturts Creek North, Birrinbubu Station, Maconochie 1753
(CANB, NT); Top Springs, Telford 11659 (CANB, DNA, M, NE, NSW, US);
Stuart Swamp, 5 km NNE of Daly Waters, Latz 13729 (CANB, DNA);
Rockhampton Downs, Government Paddock 7, Wilson N402 (CANB, K,
NT). Queensland: Camooweal Caves National Park, Nowranie Waterhole,
McDonald & Dennis 8916 (BRI, NE); Mount Eliza, 8.5 km along O’Briens
Creek Road from Mount Surprise township, Telford & Sebastian 13314 (M,
NE), our Fig. 2A , B. New South Wales: Kirramingly Nature Reserve, ca. 30
km WSW of Moree, Nano & M. Gardner NE96519 (NE); ‘Burrenda’, near
Burren Junction, Carrington CANB 310499 (CANB).
Distribution— Endemic to northern Australia from the
Hamersley Ranges, Western Australia eastwards to Rock-
hampton, Queensland, and southwards in the Murray –
Darling Basin to Burren Junction, New South Wales ( Fig. 3 ).
Habitat—Cucumis picrocarpus grows in a variety of habitats,
particularly on clay flats, such as alluvium on flood plains, but
recorded also from rocky loam on hillsides. Vegetation com-
munities include Astrebla grassland, grassy open woodland
with Eucalyptus , Bauhinia, Terminalia and riverine woodland
dominated by Eucalyptus camaldulensis Dehnh. and E. microth-
eca F. Muell.
Phenology— Flowers and fruits February–May.
Conservation Status— The species is widespread and com-
mon and is not considered at risk.
Notes— Von Mueller (1859) in the protologue described the
fruits as extremely bitter. Several specimens in CANB and
MEL were annotated “ Cucumis melo subsp. nov.” by Charles
Jeffrey in 1986. Cucumis picrocarpus is sympatric with C. melo
over much of its range ( Fig. 3 ) but no intergrading or hybrid-
ization has been observed.
Cucumis melo L., Sp. Pl. ed. 1, 1011. 1753. TYPE: SWEDEN:
Plant cultivated at Uppsala (lectotype: LINN, sheet
number 1152.8, photograph!); see discussion in A. D. J.
Meeuse (1962) .
Cucumis jucundus F. Muell., Trans. Philos. Inst. Victoria 3:
46. 1859 as ‘jucunda’. TYPE: AUSTRALIA. Northern
Territory: Victoria River, Mueller s. n . (neotype K, pho-
tograph!; isoneotype: K, photograph!), fide Kirkbride
(1993) .
Cucumis pubescens T. Mitch., J. Exped. Tropical Australia 110.
1848 nom. nud., non Willd. 1805.—TYPE: AUSTRALIA.
31 March 1848, latitude of camp, 28°38’47”S, Mitchell s. n.
(holotype: K).
Trailing or climbing annual herb, hispid on most parts,
monoecious; stems to 3 m long, 2.6 mm diam, ribbed. Tendrils
simple, to 9 cm long. Leaves: petiole 15–80 mm long; lamina
ovate in outline, 3–78 × 2.6–74 mm, unlobed or palmately shal-
lowly 3- or 5-lobed, mucronate, denticulate. Inflorescences
unisexual. Male flowers in (1–)3–5-flowered fascicles; pedi-
cels 3–13 mm long; hypanthium narrowly campanulate, 3.5–
4.3 mm long, pilose with spreading hairs; calyx lobes linear to
narrowly triangular, 2–4 mm long; corolla lobes ovate to ellip-
tic, 8–12 × 4–6 mm, mucronate, glabrous or sparsely hispid
outside, glabrous inside, yellow; stamens inserted towards
the base of the hypanthium tube; anthers one 1-thecous, two
2-thecous, sigmoid, ca. 2 mm long; connective appendage
1–2.2 mm long, lobed, papillose; disc depressed subglobose,
ca. 0.8 mm diam. Female flowers solitary; pedicels 2.8–13 mm
long; ovary ellipsoidal, 5–8 mm long, 2–3.4 mm diam, pilose
with antrorse or spreading hairs; hypanthium above constric-
tion 2.4–3 mm long; perianth similar to male; disc shortly
cylindrical, ca. 1.3 mm diam, lobed; style 1–1.4 mm long; stig-
matic branches 1.3–1.8 mm long. Fruit ellipsoidal, 20–36 mm
long and 14–24 mm in diam, with short antrorse hairs, fruit-
ing pedicel to 14 mm long. Seeds elliptic, 4.2–5 × 2–2.7 mm,
± compressed with the faces slightly convex, smooth, buff.
Representative Specimens Examined— AUSTRALIA. Western Australia:
ca. 10 km W of Mount Vernon Station Homestead near Meekatharra,
Mitchell 4713 (CANB, PERTH); Hamersley Ranges National Park, on
flats E of Mount Bruce, Trudgen 2578 (PERTH), Fig. 2C shows an image
of this specimen; Harding River bridge, Roebourne, Olsson 57 (PERTH);
ca. 100 km S of Dampier on Hamersley Iron Railway line road, Mitchell
PRP231 (CANB, PERTH); De Grey River, Burbidge 948 (PERTH). Northern
Territory: Jasper Gorge, Victoria Downs road crossing of Jasper Creek,
Telford 11655 (BRI, CANB, DNA, NE, US). South Australia: Oodnadatta,
Knight 242 (AD); Lake Eyre basin, Scrubby Camp Waterhole, Thorpe 85
(AD). Queensland: 47.9 km by road SW from Burke – Gulf Development
Road junction near Normanton, McDonald & Dennis 8872 (BRI, NE);
Mount Eliza, 8.5 km along O’Briens Creek Road from Mount Surprise
township, Telford & Sebastian 13313 (M, NE), Fig. 2D shows an image of
this collection; Long Hole, Winton Water Supply, Forster & Booth 22315
Distribution— Widespread in Australia across the tropics
and subtropics from near Meekatharra, Western Australia to
Rockhampton, Queensland, south to Lake Eyre South, South
Australia, and near Wilcannia, New South Wales ( Fig. 3 ).
F ig . 2. Morphological traits of Australian Cucumis species. A, B. Cucumis picrocarpus. A. Flowering stems. B. Developing fruit (both from Telford &
Sebastian 13314 ). C, D. C. melo . C. Flowering and fruiting stems (from Trudgen 2578 ); D. fruit (from Telford & Sebastian 13313 ). E, F. C. althaeoides . E. Male inflo-
rescence. F. Fruiting stem. (both from Copeland & Bell 4220 ). Scale bars = 1 cm. Images A, C by J. J. Bruhl; B, D by P. Sebastian; E, F by L. M. Copeland.
Habitat—Cucumis melo grows in a variety of habitats
including grasslands on cracking clays, Eucalyptus , Corymbia ,
Acacia, or Grevillea grassy woodlands on clay flats, less com-
monly on rocky slopes.
Phenology— Flowers and fruits January–May.
Conservation Status— The species is widespread and com-
mon and is not considered at risk. In Western Australia, the
species is recorded as “common all along the Ashburton
floodplain” (label data from Mitchell PRP231 ).
Notes— The description above is based on Australian
material. This taxon was treated by Kirkbride (1993) as
Cucumis melo L. subsp. agrestis (Naudin) Pangalo, a subspe-
cies thought to occur throughout Africa, Asia, and Australia.
The Australian populations appear to be indigenous as early
European explorers already found C. melo in inland Australia
before the establishment of Western settlements and gardens
there. Because the fruits are wild food of aboriginal people,
there is the possibility of anthropogenic range expansion, but
resolving whether humans account for the species’ arrival in
Australia will require a detailed phylogeographic study of
C. melo .
The aboriginal people in the Pilbara eat the fruits as a wild
food (label data from Burbidge 948 ); the local aboriginal name
is “Yindjibarndi” (label data from Olsson 57 ). In the Lake Eyre
region, South Australia, “eaten by the Aborigines when falls
off vine, rubbed in sand first”; local name “Ilcarta” (label data
from Knight 242 ).
Plants of domestic cultivars are occasionally found as
weeds around picnic areas (e.g. New South Wales: Namoi
River, Warrabah National Park, Hosking 941 ) but they do not
appear to persist.
Cucumis costatus I. Telford sp. nov. TYPE: AUSTRALIA.
Queensland: Cook District: Mt Scatterbrain, Butchers
Hill Station, near Lakeland, 15°52’ S 144°53’ E, 25 Jan.
1992, Forster 9514 (holotype: BRI!; isotypes: CNS!, M!,
MEL!, NE!).
Mukia sp. B, I. R. H. Telford, Fl. Australia 8: 186. 1982.
F ig . 3. Distributions of Cucumis picrocarpus (solid circle); C. melo (hollow circle).
Mukia racemosa I. Telford, J. D. Briggs & J. Leigh, Rare or
Threatened Australian Plants ed. 2: 44. 1995 nom. nud .
Mukia sp. (Little Annan R., B. Gray 101 ), R. J. F. Henderson (ed.),
Names & Distributions of Queensland Plants 53. 2002;
P. Bostock & A. E. Holland, Census of the Queensland
Flora 54. 2007.
Ab affini Cucumi javanico differt floribus masculis in rac-
emo digestis, fructu costato seminibusque turgidis.
Trailing or climbing herb, monoecious, hispid on most veg-
etative parts; stems to ca. 1.5 m long, ca. 1 mm diam, ribbed,
sparsely hispid, annually sprouting from a thickened peren-
nating rootstock. Tendrils simple, to 40 mm long. Leaves: peti-
oles 10–33 mm long; lamina triangular to ovate, 25–35 × 18–28
mm, cordate, scarcely lobed or shallowly 3- or 5-lobed, den-
tate, the lobes obtuse, mucronate, hispid on both surfaces with
hairs to 0.7 mm long. Inflorescences unisexual. Male flowers
in 4–11-flowered racemes 25–37 mm long; peduncles 3–30 mm
long; pedicels 3–12 mm long; hypanthium narrowly campan-
ulate, 2.6–5.3 mm long, hispid; calyx lobes 5, narrow-trian-
gular, 0.6–2 mm long; corolla lobes 5, rotate, ovate, 2.8–5.2 ×
1–2.2 mm, apex rounded, glabrous or sparsely hispid outside,
glabrous inside, yellow; stamens inserted about the middle of
the hypanthium tube; anthers one 1-thecous, two 2-thecous,
straight, ca. 2.2 mm long; connective appendage minute; disc
depressed subglobose, ca. 1.3 mm diam. Female flowers sol-
itary; peduncles 10–14 mm long, elongating in fruit; ovary
narrowly fusiform, attenuate, 6–9.5 mm long ca. 2 mm diam,
hispid with retrorse multicellular hairs; hypanthium above
constriction ca. 3 mm long; perianth similar to male; stamin-
odes absent; disc annular, ca. 1 mm diam; style ca. 1.8 mm long;
stigma 3-branched, the branches ca. 1 mm long, each bifid for
about half their length. Fruit fusiform or ellipsoidal, 16–32 mm
long, 10–12 mm diam, longitudinally 9- or 10-ribbed, sparsely
hispid, scabrid on ribs with retrorse tubercle-based hairs,
green, 15–30-seeded; pericarp thin, showing seeds when dry;
fruiting peduncle up to 40 mm long. Seeds ovate, 3.4–4 × 2.6–3
mm, pale brown, the faces convex, pitted, the margin narrow.
Representative Specimens Examined— AUSTRALIA. Queensland:
McIvor River area, 0.5 km N of Tribulation Creek, 45 km NW of Cooktown,
Wannan & Addicot 3845 (BRI); McIvor River, 5 km SW of Mt Ray, Wannan &
Lyon 3931 (BRI, CNS, NE, NSW), our Fig. 4A and B; Endeavour River,
Persieh 169 (MEL); Little Annan River crossing on the Cooktown road,
Gray 101 (CNS); 0.9 km E of the West Normanby River on the Lakeland
Downs to Cooktown road, Clarkson &. MacDonald 6747 (BRI, CANB, CNS);
northern ridge off Mount Sampson, Annan River catchment, McDonald
et al. 2086 (BRI).
Distribution— Endemic to North-Eastern Queensland
on the eastern fall of the Great Dividing Range in southern
Cape York Peninsula, the species is recorded from the drain-
age basins of the McIvor, Endeavour, Annan, and Normanby
Rivers ( Fig. 5 ).
Habitat—Cucumis costatus grows in gallery forest or euca-
lypt woodland and depauperate deciduous vine thicket on
low rocky hills, on basalt or metamorphics, to 300 m altitude.
Associated species recorded include Eucalyptus platyphylla
F. Muell ., E. leptophleba F. Muell. and Erythrophleum chloros-
tachys (F. Muell.) Baillon.
Phenology— Flowers and fruits January–April.
Conservation Status— The species was allocated category
2K in Briggs and Leigh (1995) , meaning it is a poorly known
taxon with a geographic range of less than 100 km. At the type
locality, it is recorded as “very common.” Additional recent col-
lections indicate that C. costatus is more widespread than previ-
ously thought, and we therefore do not consider it at risk.
Etymology— From Latin costatus (ribbed), in reference to
the pronounced longitudinal ribbing of the fruit.
Notes—Cucumis costatus was the first Australian Cucumis
species sequenced for nuclear and chloroplast regions and is
included in the phylogeny of Renner et al. (2007) as Cucumis sp.
HS414. It differs from the morphologically similar C. javanicus
in the male flowers being borne in racemes and costate fruits.
Cucumis queenslandicus I. Telford sp. nov. TYPE:
AUSTRALIA. Queensland: Cook District: 5 km SE of
Chillagoe, beside Burke Development Road, 18 Apr.
2006, Wannan & Beasley 4266 (holotype: BRI!; isotypes:
CANB!, CNS!, M!, NE!).
Ab affini Cucumi costato differt foliis non profunde vel pro-
funde lobatis, floribus masculis fasciculatis vel racemosis et
fructu leviter costato.
Trailing or climbing perennial herb, monoecious, hispid
on most vegetative parts; stems to 70 cm long, ribbed, annu-
ally sprouting from a thickened, perennating, ± cylindrical
rootstock. Tendrils simple, to 60 mm long. Leaves: petioles
(6–)14–40 mm long; lamina subtriangular to broadly ovate in
outline, 9.5–44 × 11.5–42 mm, cordate, shallowly or deeply 3-
or 5-lobed, when deeply lobed the lobes sinuately lobed, den-
tate, obtuse, mucronate, hispid on both surfaces with hairs
to 0.2 mm long. Inflorescences unisexual. Male flowers in
2–5-flowered fascicles or racemes with peduncles to 12 mm
long; pedicels 1.3–13 mm long; hypanthium narrowly cam-
panulate, 2.8–4.2 mm long; calyx lobes 5, linear, 0.8–1.3 mm
long; corolla lobes 5, obovate, 3.4–4.2 × 2–2.5 mm, rounded,
glabrous or sparsely hispid outside, glabrous inside, yellow;
stamens 3, inserted about the middle of the hypanthium tube;
anthers one 1-thecous, two 2-thecous, straight, ca. 2 mm long;
connective appendages minute; disc depressed globose, ca.
1.2 mm diam. Female flowers solitary; peduncle 5–18 mm
long, scarcely elongating in fruit; ovary narrowly fusiform,
ca. 5.5 mm long, ca. 1.2 mm diam, attenuate, densely hispid
with retrorse, multicellular hairs; hypanthium above con-
striction ca. 6 mm long; perianth similar to male; disc annu-
lar, undulate, ca. 0.5 mm diam; style ca. 1.3 mm long; stigma
capitate, ca. 2.5 mm long; staminodes absent. Fruit ovoid–
fusiform, 15–23 mm long, 9–10 mm diam, slightly 10-ribbed,
hispid, 10–20-seeded; pericarp thin, showing seeds when dry;
fruiting pedicel 7–16 mm long. Seeds ovate, 4.5–5 × 3.8–4.2
mm, pale brown, the faces convex, pitted, the margin narrow.
Figure 4C shows an image of the type collection.
Representative Specimens Examined— AUSTRALIA: Queensland:
Cook District: Metal Hills section, Chillagoe − Mungana Caves National
Park, McDonald 8814 (BRI, NE), Fig. 4D shows an image of this collection;
Mt Eliza, 8 km NW of Mt Surprise, Forster & Bean 12812 (BRI); Mt Eliza,
8.5 km along O’Briens Creek Road from Mount Surprise township, Telford &
Sebastian 13316 (BRI, CNS, M, NE); Undara National Park, Quartz Blow
Lookout, McDonald 3662 (BRI); Newcastle Range, eastern flank, W of
Mount Surprise, McDonald 3393 (BRI); near Copperfield River Dam,
Kidston, Bean & Forster 7443 (BRI).
Distribution—Cucumis queenslandicus is endemic to North-
Eastern Queensland on the western fall of the Great Dividing
Range where it is known from near Chillagoe and from
Undara Volcanic National Park to the Newcastle Range and
south to the Copperfield River ( Fig. 5 ).
Habitat— The species grows in Eucalyptus woodland on
granite (Undara Volcanic National Park), the granite/lime-
stone contact zone (Chillagoe), and deciduous vine thickets
on granite hills (Mount Eliza and Undara Volcanic National
Park) at 400–550 m altitude. Associated species recorded
F ig . 4. Morphological traits of Australian Cucumis species. A, B. Cucumis costatus . A. Male inflorescences. B. Fruit (both from Wannan & Lyon 3931 ).
C, D. C. queenslandicus . C. Male inflorescences (from Wannan & Beasley 4266 ). D. Fruit (from McDonald 8814 ). E, F. C. umbellatus . E. Flowering stems with
developing fruit (from Sebastian 14 ). F. Fruit (from Wannan, Wardrop & Lane 5728 ). Scale bars = 1 cm. Images A–C, F by B. S. Wannan; D by K. R. McDonald;
E by J. J. Bruhl.
include Eucalyptus cullenii Cambage, E. miniata Schauer,
Erythrophloeum chlorostachys , Brachychiton chillagoensis Guymer
and B. albidus Guymer.
Phenology— Flowers and fruits January–June, mainly
Conservation Status—Cucumis queenslandicus is wide-
spread with several recent collections in the Newcastle Range
and Mount Surprise areas and is not considered at risk. It
is conserved in Chillagoe − Mungana Caves and Undara
Volcanic National Parks.
Etymology— Named for the state which includes the area
of endemism of the species in north-eastern Queensland.
Notes— Molecular data ( Fig. 1 ) corroborate an affinity
to C. costatus and C. umbellatus . Cucumis costatus differs in
having male flowers in more elongate racemes and strongly-
ribbed fruit, C. umbellatus in umbellate male inflorescences
and small unribbed fruit.
Cucumis umbellatus I. Telford sp. nov. TYPE: AUSTRALIA.
Northern Territory: Darwin and Gulf: Kakadu National
Park, Ubirr, Apr. 2009, Sebastian 14 (holotype: CANB!;
isotypes, DNA!, K!, M!, MO!, NE!, PERTH!).
Cucumis sp. Gunlom ( J. L. McKean 864b ), Council of Heads
of Australian Herbaria, Australian Plant Census. 2008
( )
Mukia sp. A, I. R. H. Telford, Fl. Australia 8: 184. 1982; J. R.
Wheeler, Fl. Kimberley Region 252. 1992.
Ab affini Cucumi costatus differt floribus masculis umbella-
tis, fructibus longissime pedicellatis subglobosis vel ellipsoi-
dalibus et seminibus paucioribus.
Trailing or climbing annual herb, monoecious, most vegeta-
tive parts sparsely hispid; stems to 2 m long, to 1 mm diam,
ribbed. Tendrils simple, to 2 cm long. Leaves: petiole 8–35
mm long; lamina broadly ovate in outline, 10–40 × 14–45 mm,
cordate, shallowly 3- or 5-lobed, dentate, the lobes obtuse,
mucronate, sparsely hispid on both surfaces with hairs to
1.8 mm long. Inflorescences unisexual. Male flowers in 12–20-
flowered umbels; peduncles 10–55 mm long; pedicels 3–13 mm
long; hypanthium campanulate, 1.6–2.2 mm long, sparsely
hispidulous; calyx lobes linear, 0.5–0.8 mm long, sparsely
hispidulous; corolla lobes ovate, 2.6–4 × 1.6–2 mm, obtuse,
mucronate, glabrous or sparsely hispidulous outside, glabrous
inside, yellow; stamens inserted about the middle of the hypan-
thium tube; anthers one 1-thecous, two 2-thecous, straight, ca.
1.2 mm long; connective appendages minute; disc depressed
subglobose, ca 0.7 mm diam. Female flowers solitary, rarely
paired in axils; peduncle 5–25 mm long, greatly elongating
in fruit; ovary subglobose to fusiform, ca. 0.8–1.2 mm diam,
hispid with retrorse hairs; hypanthium above the constric-
tion narrowly campanulate, 1.4–1.8 mm long; perianth simi-
lar to male; disc annular, ca. 0.6 mm diam; style ca. 0.8 mm
long; stigmatic branches 0.8–1 mm long. Fruit subglobose, 4–7
mm diam, hispid with tubercle-based hairs, ripening bright
orange, with 1 or 2, rarely 3 seeds; fruiting peduncle 50–115
mm long. Seeds ovate, 4.5–5 × 2.8–3.4 mm, buff, the faces con-
vex, deeply pitted, the margin narrow. Figure 4E shows an
image of the type specimen.
Representative Specimens Examined— AUSTRALIA. Western
Australia: 15 km N of Mount Disaster, Keighery 10601 (PERTH); Prince
Regent River, Wannan et al. 5728 (BRI, PERTH), Fig. 4F shows an image
of this collection; island inside Yule Entrance, Walcott Inlet, Mitchell 3531
(CANB, NE, PERTH); Boongarree Is., 18.3 km NE of Mount Knight,
Keighery 10695 (CANB, PERTH); mouth of Glenelg River, 20.5 km NW of
Mount French, Keighery 10689 (PERTH); tributary of Camp Creek, ca. 12
km SW of CRA Mining Camp, Mitchell Plateau, Kenneally 7909 (PERTH);
Carson Escarpment, 2.5 km N of Face Point, Keighery 10667 (PERTH);
Cambridge Gulf, 1887, Wright (MEL). Northern Territory: Obiri [Ubirr]
Rock track, 4 km NW of Cahills Crossing, East Alligator River, Telford &
Wrigley 7631 (CANB, DNA, K, NE, PERTH);1 mile SW of Cannon Hill,
Martenz & Schodde AE 647 (BRI, CANB, DNA, NT); Kakadu National
Park, Gubarra, Sebastian 15 (BRI, CANB, DNA, L, M, NE, US); Baroalba
F ig . 5. Distributions of Cucumis costatus (circle); C. queenslandicus (triangle) C. umbellatus (square).
Creek, ca. 1 km E of gorge entrance, Cunliffe s. n . (CANB, DNA, NSW, OSS,
UNSW); Little Nourlangie Rock, Telford & Wrigley 7814 (BISH, CANB);
headwaters of Liverpool River, Craven & Wightman 8363 (CANB).
Distribution—Cucumis umbellatus is endemic to northern
Australia, occurring in the western and northern Kimberley
region and disjunctly in the Northern Territory along the
northwestern escarpment of Arnhem Land ( Fig. 5 ).
Habitat— The species grows in sandy soils on sand-
stone, usually on rock outcrops or talus slopes or in rocky
watercourses. The species occurs in vine thickets together
with Allosyncarpia ternata S. T. Blake forest, in low open wood-
land with Xanthostemon paradoxus F. Muell. and Terminalia had-
leyana W. Fitzg., and in open shrubland with Triodia species.
Phenology— Flowering and fruiting February–May.
Conservation status—Cucumis umbellatus is widespread in
the Kimberley region, where it is conserved in Drysdale River
National Park, and it is also common in Kakadu National
Park. The species is therefore not considered at risk.
Etymology— From the Latin umbella (parasol), in reference
to the inflorescences of the male flowers.
Notes— The peduncle of the female flower elongates up to
115 mm as the fruit develops. This elongation may push the
ripening fruit into the protection of crevices in the rocky slopes
of its habitat. In the African Cucumis humifructus , the fruits are
geocarpic; buried by the elongating pedicel ( Meeuse, 1962 ).
Although separated by a disjunction of some 500 km, no mor-
phological differences between Kimberley and Top End pop-
ulations are apparent.
Cucumis argenteus (Domin) P. Sebastian & I. Telford, comb.
nov. Melothria argentea Domin, Biblioth. Bot. 89: 635, Fig.
196. 1929. TYPE: AUSTRALIA. Queensland: Flinders
River near the town Hughenden, Feb. 1910, K. Domin
8716 (holotype: PR, photograph!)
Melothria celebica var. villosior Cogn., Bull. Acad. Roy. Sci.
Belgique ser. 3, 14: 357. 1887. TYPE: AUSTRALIA. Gulf
of Carpentaria, Mueller s. n. (holotype: BR photograph!;
isotype: MEL!).
Mukia maderaspatana auct. non (L.) M. Roem.: I. R. H. Telford,
Fl. Australia 8: 183, 185 Fig. 40E, G. 1982.
Trailing or climbing herb, monoecious, most vegetative
parts pilose or densely hispid; stems to 3 m long, to 2 mm
diam, ribbed. Tendrils simple, to 9 cm long. Leaves: sub-
sessile or petiole to 28 mm long; lamina triangular, lanceolate
or broadly ovate in outline, 24–93 × 14–87 mm, cordate, acute
or obtuse, mucronate, shallowly 3- or 5- lobed, denticulate,
densely scabrid or villous with hairs to 1.2 mm long on both
surfaces. Inflorescences unisexual. Male flowers in 2–5-flow-
ered fascicles; pedicels to 7 mm long; hypanthium campanu-
late, 3–3.5 mm long, pilose or hispid; calyx lobes linear, ca. 1
mm long, pilose or hispid; corolla lobes ovate, 3.4–3.6 × 2.5–
2.7 mm, rounded, sparsely pilose or puberulous outside, the
hairs denser apically, puberulous inside, pale yellow; stamens
inserted about the middle of the hypanthium tube; anthers
one 1-thecous, two 2-thecous, straight, ca. 2 mm long; connec-
tive appendages minute; disc depressed subglobose, 0.7–1.3
mm diam. Female flowers 1 or 2 per axil, sometimes coaxil-
lary with several males; subsessile or pedicels to 2 mm long;
ovary subglobose, ca. 1.5 mm diam, white pilose with antrorse
hairs; hypanthium above the constriction narrowly campanu-
late, 2.4–3 mm long; perianth similar to male; disc annular,
1–1.5 mm diam; style ca. 1.8 mm long; stigmatic branches ca.
1.8 mm long. Fruits globose, 6.4–8 mm diam, pale green with
darker longitudinal markings, at maturity sparsely or densely
pilose, red, with 5 seeds; fruiting pedicel to 2.5 mm long. Seeds
ellipsoidal, 4.5–4.8 × 2.8–3.3 mm, the faces convex, smooth,
lacking a thickened margin, grey-black or pale brown.
Representative Specimens Examined— AUSTRALIA. Western Australia:
246 km from Broome along Great Northern Highway towards Port
Hedland, Telford & Butler 6028 (BISH, CANB, K, MO, PERTH); Wolf Creek
Crater, George 15338 (NT, PERTH). Northern Territory: Petermann Ranges,
Lasseters Cave, Henshall 3450 (AD, DNA, NT); Kings Canyon National
Park, Bruhl & Quinn 2162 (L, NE); Ilparpa Road, Alice Springs, Albrecht
10202 (CANB, NT), Fig. 6B shows an image of this specimen; West Island,
Sir Edward Pellew Group, Braithwaite 3290 (CANB). South Australia:
Lake Eyre region, Peake Creek, Conrick 724 (AD, CANB). Queensland:
Riversleigh archaeological site D, Australian Geographic information area,
Barnsley 1656 (BRI, CANB, NE), Fig. 6A shows an image of this specimen;
20.2 km by road S of Musselbrook Mining Camp on road to Camooweal,
Thomas & Johnson s. n. (A, AD, BRI, DNA, K, NE); 3 km from Mount Isa,
Ollerenshaw & Kratzing 1161 (BRI, CANB, L).
Distribution— Endemic to Australia, Cucumis argenteus
occurs widely from Ninety Mile Beach, Western Australia,
southwards to the Flinders Ranges, South Australia, east-
wards to Winton, Queensland ( Fig. 7 ).
Habitat—Cucumis argenteus grows in a range of habitats
from sand plains, loamy flats, rocky hillsides and limestone
ridges to tussock grassland, Triodia hummock grassland, open
shrublands and eucalypt or Acacia woodland.
Phenology— Flowers and fruits throughout the year, pos-
sibly in response to rainfall, but mostly between April and
August in tropical areas.
Notes— Three collections by K. Domin of Melothria argentea
from the Flinders River near Hughenden, Queensland, are
held in PR, numbered Domin 8715, 1816 , and 8717 . Although
no collector’s number is given in the protologue, the image is
of K. Domin 1816, with the caption “ Melothria argentea Dom.
nach dem Originalexemplare von Flinders River frequenter.”
The species differs from C. rumphianus in its smaller fruit
and seeds. Being widely distributed, the species is accord-
ingly variable in leaf size and indumentum length and den-
sity. Collections from the Barkly Tableland and its northern
slopes, Queensland, are densely white hairy; plants further
inland are scabrid. Specimens with the dense, whitish indu-
mentum only on the lower leaf surface have been collected
(Braithwaite 3290 , Ollerenshaw & Kratzing 1161 ). Further study
may lead to the recognition of two subspecies.
Cucumis althaeoides (Ser.) P. Sebastian & I. Telford, comb.
nov. Bryonia althaeoides Ser. in DC. Prodr. 3: 306. 1828;
Mukia althaeoides (Ser.) M. Roem. Syn. Monogr. 2: 47.
1846; Melothria althaeoides (Ser.) Nakai, J. Jap. Bot. 14: 127
1938 (as “ altaeoides ”). TYPE: TIMOR. leg. ign . (holotype:
G-DC, photograph!).
Mukia maderaspatana auct. non (L.) M. Roem.: I. R. H. Telford,
Fl. Australia 8: 183, 185 Fig. 40F. 1982; W. J. J. O. de Wilde
& B. E. E. Duyfjes, Thai For. Bull. (Bot.) 34: 43.2006 p. p.
Trailing or climbing perennial herb, monoecious, most veg-
etative parts hispid, sometimes pilose; stems to 3 m long, to
1.6 mm diam, ribbed, annually sprouting from a perennating
rootstock. Tendrils simple, to 15 cm long. Leaves: subsessile
or petiole to 38 mm long; lamina ovate or broadly lanceo-
late in outline, sometimes hastate, 24–75 × 18–70 mm, cor-
date, unlobed or shallowly 3-lobed, rarely 5-lobed, obtuse
or acute, mucronate. Inflorescences unisexual. Male flow-
ers in 3–10(–15)-flowered fascicles, sometimes in racemes
with peduncles to 3 mm long; pedicels to 7 mm long; hypan-
thium narrowly campanulate, 3–3.4 mm long, hispid or pilose
outside; calyx lobes linear, 0.8–1.4 mm long, hispid or pilose;
corolla lobes ovate, 3.8–4.2 × 2.6–2.8 mm, rounded, mucronate,
sparsely hispid outside, the hairs denser apically, glabrous or
hispid along the veins inside, hispid around the mouth of the
hypanthium tube, bright yellow; stamens inserted about the
middle of the hypanthium tube; filaments ca. 0.4 mm long;
anthers one 1-thecous, two 2-thecous, straight, 1.5–2 mm
long; connective appendages minute; disc depressed globose,
0.8–1.4 mm diam. Female flowers 1 or 2, rarely 3 or 4 per
axil; pedicels to 2 mm long; ovary ellipsoidal, ca. 3 mm long,
pilose with antrorse hairs; hypanthium above the constric-
tion narrowly campanulate, ca. 3 mm long; perianth similar
to male; disc annular, 1–1.2 mm diam; style ca.1.5 mm long;
stigmatic lobes 3, ca. 1.3 mm long. Fruit globose, 8–18 mm
diam, pale green with darker longitudinal markings, at matu-
rity sparsely pilose, red, with 9–20(–25) seeds; fruiting pedi-
cel to 6 mm long. Seeds ovate, 3.8–4.5 × 2.3–2.8 mm, buff, the
faces convex, verrucose, the margin thickened, raised.
Representative Specimens Examined— INDONESIA: Timor, Zippel
107 (L). PAPUA NEW GUINEA: near Matapaili village, Darbyshire 696
F ig . 6. Morphological traits of Australian Cucumis species. A, B. Cucumis argenteus . A. Typical specimen from Barkly Tableland, Queensland ( Barnsley
1656 ). B. Typical specimen from Central Australia ( Albrecht 10202 ). C–F. C. variabilis . C. Typical specimen from southern Pilbara, Western Australia (from
Byrne 2809 ). D. Variant from North West Cape, Western Australia (from Wajon 473 ). E. Variant from Barrow Island, Western Australia (from White 87 ).
F. Pilbara variant (from Payne PRP 1864 ). Scale bars = 1 cm. Images by J. J. Bruhl.
(A, CANB, L, LAE); S coast near Hood Bay, Paijmans 775 (CANB); SE side
of Little Mount Lawes, ca. 16 miles N of Port Moresby, Pullen 6804 (A,
BRI, CANB, L, LAE, K, TNS). AUSTRALIA: Western Australia: Glycosmis
Bay, Mitchell 7726 (PERTH). Northern Territory: Humpty Doo, McKee 8320
(DNA); 13 km SW of Cape Arnhem, Brennan 2576 (DNA). Queensland:
Lizard Island, Mangrove Beach, Batianoff & Tarte 12083 (A, BRI, DNA, K, LE);
McIvor River, “Mount Ray”, 1.5 km NW of Mount Ray, Telford & Sebastian
13308 (BRI, CNS, L, M, NE); Bucasia Beach, 13 km N of Mackay, Clarke
NE 81288 (CANB, M, NE); Tannum Sands, 15 km SE of Gladstone, Telford
5499 (CANB). New South Wales: Oxley Wild Rivers National Park, Aspley
River, 300 m downstream from Rusdens Creek, Copeland & Bell 4220
(CANB, M, NE, NSW), our Fig. 2E and F .
Distribution—Cucumis althaeoides occurs in coastal and sub-
coastal Timor, southern New Guinea and northern Australia,
from the Kimberley region in Western Australia eastwards
through the Northern Territory, Queensland and south to
near Walcha, New South Wales ( Fig. 7 ).
Nakai (1938) cites collections from the Ryukyu Islands
and Taiwan, and states that the species also occurs on the
Philippines, Borneo, New Guinea, and Timor (besides Australia).
As yet, we have not studied specimens from north of Timor.
Habitat— The species grows on coastal sands or riverine
alluvium in herbfields and Casuarina or Eucalyptus woodland.
On coralline beach sands of eastern Queensland, the species
is recorded as growing under Casuarina equisetifolia L. and
Pandanus tectorius Parkinson ex J. P. du Roi.
Phenology— Flowers and fruits April–August.
Conservation Status— Widespread and common, the spe-
cies is not considered at risk.
Notes—Cucumis althaeoides differs from other Australian
species in its verrucose seeds with thickened margins. Why
Nakai (1938) treated the species under Melothria is unclear
to us. The species shows considerable variation in leaf
morphology over its range, notably in petiole length and
degree of lobing of the lamina. In southern Papua New
Guinea and northeastern Queensland, plants bearing larger,
Fig . 7. Distributions of Cucumis althaeoides (solid circle); C. argenteus (hollow circle); C. variabilis (square).
many-seeded fruits (up to 1.8 mm diam long, with 16–20
seeds) occur and perhaps warrant formal recognition at the
rank of subspecies.
Besides C. althaeoides , a second species [ A. R. Insani SAN 10
(L), collector ignotus 1305 (L), H. Raap 499 (L), the latter in our
Fig. 1 ] occurs on Timor and westwards to at least Java. Based
on the molecular data, this still unnamed species is sister to C.
althaeoides and C. variabilis .
Cucumis variabilis P. Sebastian & I. Telford sp. nov. TYPE:
AUSTRALIA. Western Australia: 46.5 km along North
West Coastal Highway from Ashburton River bridge
towards Carnarvon, 25 Apr. 1992, Telford 11578 (holo:
PERTH!; iso: BRI!, CANB!, K!).
Mukia maderaspatana auct. non (L.) M. Roem.: I. R. H. Telford,
Fl. Australia 8: 183. 1982.
Ab affini Cucumi althaeoidi differt foliis non profunde vel
profunde lobatis, lobis plerumque angustissimis, semini-
busque laevibus margine carentibus.
Trailing or climbing herb, monoecious, most vegetative
parts hispid or pilose; stems to 2 m long, to 1.8 mm diam,
ribbed. Tendrils simple, to 13 cm long. Leaves: petiole 2.4–18
mm long; lamina triangular, lanceolate or ovate in outline,
often hastate or sagittate, 16–102 × 14–55 mm, cordate, acum-
inate, mucronate, shallowly to deeply 3- or 5-lobed, usually
sparsely serrate, on deeply lobed leaves, the lobes usually
narrow, linear. Inflorescences mostly unisexual. Male flowers
in 5–14-flowered fascicles, sometimes in racemes on pedun-
cles to 4 mm long; pedicels 3–7.5(–12) mm long, at first pilose;
hypanthium campanulate, 3.5–4.2 mm long; calyx lobes lin-
ear, 1–1.3 mm long, hispidulous outside; corolla lobes ovate,
4.6–5 × 3–3.4 mm, obtuse, mucronate, hispidulous outside,
glabrous inside, yellow; stamens inserted about the middle of
the hypanthium tube; filaments 0.6–1.2 mm long; anthers one
1-thecous, two 2-thecous, straight, 2.2–2.6 mm long; connective
narrow; appendages minute; disc depressed subglobose, ca. 1.2
mm diam. Female flowers in 3–7-flowered fascicles; subsessile
or pedicels to 8 mm long; ovary subglobose, ca. 1.8 diam, his-
pidulous with antrorse hairs; hypanthium above the constric-
tion narrowly campanulate, ca. 4 mm long; perianth similar to
male; disc annular, ca. 1.4 mm diam, style ca. 2 mm long; stig-
matic branches 0.8–1.2 mm long. Fruit globose, 5–8 mm diam,
at maturity glabrescent, ripening orange-red, with 10–15 seeds;
fruiting peduncle to 8.5 mm long. Seeds ovate, 5.6–6.5 × 3–3.4
mm, the faces convex, smooth, minutely verrucose in two lines
around the nonprojecting margin, grey or pale brown.
Representative Specimens Examined— AUSTRALIA. Western
Australia: Barrow Island, 75 m E of turnoff to R33, White 87 (PERTH),
Fig. 6E shows an image of this specimen; Yardie Creek Road, Cape Range
National Park, near Exmouth, Wajon 473 (PERTH), Fig. 6D shows an image
of this specimen; 3 miles N of Exmouth, Allan 469 (CANB, MEL, PERTH);
Carawine Pool, ca. 140 km SE of Shay Gap, Newbey 10312 (PERTH); Cape
Range, road to nos 3 and 4 wells, George 2470 (PERTH); ca. 25 km N of
Marble Bar on small track W of Shay Gap road, Mitchell PRP905 (CANB,
PERTH); Shellborough, 40 km NNW of Goldsworthy, Telford & Butler
6476 (CANB, PERTH); coast NW of Ilgarene Well, De Gray Station, Payne
PRP1864 (PERTH), Fig. 6F shows an image of this specimen; ca. 23.4
km WNW of De Grey Homestead, adjacent to Freshwater Bore, Mitchell
PRP868 (CANB, PERTH); Roy Hill, Byrne 2809 (PERTH), Fig. 6C shows an
image of this specimen; Barrabiddy Creek, Ashby 2944 (PERTH); 45 km E
of Carnarvon, Wilson 8389 (CANB, L, PERTH); 10 km ENE of West Angela
Hill, Hamersley Ranges, Trudgen 15877 (PERTH); 11.6 km WSW of Mount
Sabine, Millstream-Chichester National Park, Hamersley Ranges, Weston
20385 (PERTH).
Distribution—Cucumis variabilis is endemic to Western
Australia, where it ranges from NE of Onslow to the Cape
Range and Barrow Island, southwards to Carnarvon and the
Ashburton River and eastwards to NE of Nullagine ( Fig. 7).
Habitat— The species grows on calcareous or siliceous
sand and on rocky ridges and slopes in Eucalyptus low
open woodland, Acacia shrubland and Triodia hummock
Phenology— Flowers and fruits May–October.
Conservation Status—Cucumis variabilis is widespread
and not considered at risk. It is conserved in Cape Range,
Hamersley Ranges, and Millstream-Chichester National
Etymology— Latin variabilis (varying), in reference to the
vegetative variability of the species.
Notes—Cucumis variabilis has been illustrated in Telford
(1982 : Figs. 40G, 40E), both times as Mukia maderaspatana, show-
ing the extremes of leaf variability. The species shows two lines
of variation: leaf morphology and indumentum. More south-
erly populations inland from Carnarvon bear larger leaves with
a scattered, scabrid indumentum. Populations from North West
Cape have a similar indumentum but narrower leaf segments
with extreme reduction shown in the Barrow Island popula-
tion. This island population has been listed as a putative new
species, “ Cucumis sp. Barrow Island” ( D. W. Goodall 1264 ), in
the Western Australian “FloraBase” ( http://florabase.calm. ). In the Pilbara, plants bear thicker leaves with a
dense, white, villous indumentum similar to Barkly Tableland
populations of C. argenteus. These variants may eventually
deserve formal taxonomic recognition as subspecies.
Acknowledgments. We thank the Associate Editor Mark Simmons
and two extremely helpful reviewers for numerous corrections; the direc-
tors at BRI, CANB, DNA, MEL, PERTH and QRS (now CNS) for the loan
of material, with special thanks to Donna Lewis (DNA) and Darren Crayn
(CNS) for assistance with field work; directors at E, B, G-DC, MEL and PR
for images of specimens; Australian National Parks and Wildlife Service
for permission to collect in Kakadu National Park; Keith McDonald and
Bruce Wannan (Department of Environment and Resource Management,
Queensland) for additional collections and images of plants in their natu-
ral habitats. Werner Greuter, Palermo, provided the Latin diagnoses. Cate
McGregor and Kirsten Knox helped with map graphics. The School of
Environmental and Rural Science (UNE) provided funding for IRT.
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The Cucurbitaceae family contains about 1000 species in 96 genera. Representatives of all genera (except the extinct Khmeriosicyos) and a large percentage of the species have been sequenced for the ribosomal RNA transcribed spacer regions and variable regions of the plastid and mitochondrial genome. These data have allowed to infer evolutionary relationships in the family. The major phy-logenetic structure of the family is now clear, and this chapter includes an up-to-date phylogenetic scheme with the placement of all genera. The Cucurbitaceae clade originated in mainland Southeast Asia sometime in the Late Cretaceous, and the five deepest evolutionary divergences in the family all date to the Late Cretaceous, 70–80 Ma. Two of these ancient clades, the Gomphogyneae and Actinostemma, are now almost restricted to Asia. A third ancient group, the Triceratieae, is mainly Neotropical, except one African genus; other clades and tribes are more widespread. The economically most important genera are concentrated in the Cucurbiteae and Benincaseae, and species of Cucumis and Citrullus, with well-annotated genomes, therefore have largely comparable (homologous) linkage groups. In contrast to the relatively good data on the family's phylogeny, data on its ecology, physiology and morphological evolution are scarce and collection and study of wild species, many of them in threatened habitats is much needed.
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The Cucurbitaceae genus Austrobryonia, with four species endemic to Australia, is described, illustrated, and placed in a phylogenetic context based on molecular and morphological data. In the Flora of australia (Telford 1982), all four species were provisionally included in Mukia, but not formally described. Austrobryonia argillicola, A. centralis, A. micrantha and A. pilbarensis are adapted to arid central regions of Australia. All species are known from 7-27 localities, and their distributional ranges are allopatric. A phylogenetic analysis of plastid and nuclear DNA sequences that included all four species in a family-wide context revealed that Austrobryonia is the closest living relative to a Eurasian and Mediterranean clade consisting of Bryonia and Ecballium. An rbcL molecular clock, calibrated with Cucurbitaceae fossils, dates this rare biogeographic disjunction to minimally 42 my ago (with an error of ca +-25%), while the crown group of Austrobryonia may be about 8 my old.
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Recent molecular studies have revealed that the genus Cucumis in its current circumscription is paraphyletic. To become monophyletic, Cucumis must include five genera of the subtribe cucumerinae, namely Cucumella, Dicoelospermum, Mukia, Myrmecosicyos, and Oreosyce, already regarded as closely related by earlier workers based on morphology. The 19 species in the five genera are here formally transferred to Cucumis, resulting in 14 new combinations, two changes in status, and three new names (Cucumis indicus, C. kirkbrideana, and C. oreosyce). A description of the genus and a key to its 52 species are given, and the subgenera and sections are recircumscribed to reflect monophyletic groups. Subgenus Humifructus, with a haploid chromosome number of 12, contains only C. humifructus and C. hirsutus, while subgenus Cucumis, with n = 12 or n = 7, contains the remainder of the species. The latter is further divided into the sections Aculeatosi, Cucumella, Cucumis, Metuliferi, and Sagittati.
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Melon, Cucumis melo, and cucumber, C. sativus, are among the most widely cultivated crops worldwide. Cucumis, as traditionally conceived, is geographically centered in Africa, with C. sativus and C. hystrix thought to be the only Cucumis species in Asia. This taxonomy forms the basis for all ongoing Cucumis breeding and genomics efforts. We tested relationships among Cucumis and related genera based on DNA sequences from chloroplast gene, intron, and spacer regions (rbcL, matK, rpl20-rps12, trnL, and trnL-F), adding nuclear internal transcribed spacer sequences to resolve relationships within Cucumis. Analyses of combined chloroplast sequences (4,375 aligned nucleotides) for 123 of the 130 genera of Cucurbitaceae indicate that the genera Cucumella, Dicaelospermum, Mukia, Myrmecosicyos, and Oreosyce are embedded within Cucumis. Phylogenetic trees from nuclear sequences for these taxa are congruent, and the combined data yield a well-supported phylogeny. The nesting of the five genera in Cucumis greatly changes the natural geographic range of the genus, extending it throughout the Malesian region and into Australia. The closest relative of Cucumis is Muellerargia, with one species in Australia and Indonesia, the other in Madagascar. Cucumber and its sister species, C. hystrix, are nested among Australian, Malaysian, and Western Indian species placed in Mukia or Dicaelospermum and in one case not yet formally described. Cucumis melo is sister to this Australian/Asian clade, rather than being close to African species as previously thought. Molecular clocks indicate that the deepest divergences in Cucumis, including the split between C. melo and its Australian/Asian sister clade, go back to the mid-Eocene. Based on congruent nuclear and chloroplast phylogenies we conclude that Cucumis comprises an old Australian/Asian component that was heretofore unsuspected. Cucumis sativus evolved within this Australian/Asian clade and is phylogenetically far more distant from C. melo than implied by the current morphological classification.
The genus Mukia Arn. has been taxonomically revised. Mukia maderaspatana (L.) M. Roem. var. gracilis Kurz has been raised to specific rank, Mukia gracilis (Kurz) W.J. de Wilde & Duyfjes. The only species of the Indian endemic genus Dicoelospermum C.B. Clarke has been combined as Mukia ritchiei (C.B. Clarke) W.J. de Wilde & Duyfjes. The correct name for the eastern Malesian Mukia celebica appeared to be Mukia rumphiana (Scheff.) W.J. de Wilde & Duyfjes, of which one new subspecies is also described, Mukia rumphiana (Scheff.) W.J. de Wilde & Duyfjes subsp. tomentosa W.J. de Wilde & Duyfjes. A key to the species and descriptions of the Thai and Malesian species are presented.
The general maximum likelihood approach to the statistical estimation of phylogenies is outlined, for data in which there are a number of discrete states for each character. The details of the maximum likelihood method will depend on the details of the probabilistic model of evolution assumed. There are a very large number of possible models of evolution. For a few of the simpler models, the calculation of the likelihood of an evolutionary tree is outlined. For these models, the maximum likelihood tree will be the same as the “most parsimonious” (or minimum-steps) tree if the probability of change during the evolution of the group is assumed a priori to be very small. However, most sets of data require too many assumed state changes per character to be compatible with this assumption. Farris (1973) has argued that maximum likelihood and parsimony methods are identical under a much less restrictive set of assumptions. It is argued that the present methods are preferable to his, and a counterexample to his argument is presented. An algorithm which enables rapid calculation of the likelihood of a phylogeny is described.
The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data, In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
Recent studies show that the genus Cucumis L. in its current sense is paraphyletic, with Cucumella Chiovenda, Dicaelospermum C. B. Clarke, Mukia Arnott, Myrmecosicyos C. Jeffrey, and Oreosyce Hooker f. nested within it. A proposal to expand Cucumis to include these nested genera has therefore been made. In this paper the nomenclatural changes that are needed to accommodate the currently recognized taxa of the nested genera in Cucumis are made. The following new combinations and new names are proposed: Cucumis aetheocarpus (C. Jeffrey) Ghebretinsae & Thulin, C. bryoniifolius (Merxmüller) Ghebretinsae & Thulin, C. cinereus (Cogniaux) Ghebretinsae & Thulin, C. clavipetiolatus (J. H. Kirkbride) Ghebretinsae & Thulin, C. engleri (Gilg) Ghebretinsae & Thulin, C. gracilis (W. S. Kurz) Ghebretinsae & Thulin, the new name C. indicus Ghebretinsae & Thulin, C. javanicus (Miquel) Ghebretinsae & Thulin, C. kelleri (Cogniaux) Ghebretinsae & Thulin, the new name C. kirkbridei Ghebretinsae & Thulin, C. leiospermus (Wight & Arnott) Ghebretinsae & Thulin, C. messorius (C. Jeffrey) Ghebretinsae & Thulin, C. reticulatus (A. Fernandes & R. Fernandes) Ghebretinsae & Thulin, C. ritchiei (C. B. Clarke) Ghebretinsae & Thulin, and C. silentvalleyi (Manilal, T. Sabu & P. J. Mathew) Ghebretinsae & Thulin.