Solanum watneyi, a new bush tomato species from the Northern Territory, Australia named for Mark Watney of the book and film “The Martian”

Abstract A new species of andromonoecious Solanum from the Australian “bush tomato clade” of Solanum subgenus Leptostemonum is described. Solanum watneyi Martine & Frawley, sp. nov. is closely allied with Solanum eburneum, and is sympatric with it in parts of its range in the Northern Territory. The new species has been recognized as a variant of Solanum eburneum for decades, at times being referred to by local botanists as Solanum sp. “Bullita” because of its relative abundance in the vicinity of the Bullita Station area of Judbarra/Gregory National Park. Morphometric analyses show that Solanum watneyi differs statistically from Solanum eburneum in several key reproductive and vegetative characters and field observations suggest that the two sister species may represent a case of edaphic speciation. We provide morphometric evidence for the novelty of Solanum watneyi, a complete description, and cite specimens for both species.


Introduction
Since David Symon's monograph of Australian Solanum L. in 1981, numerous additional species and morphospecies have been described for the continent. Th is has been notably true for a set of "spiny solanums" from northern Australia defi ned by Symon (1981) as belonging to Solanum subgenus Leptostemonum Bitter section Melongena Bitter (e.g. Bean and Albrecht 2008, Barrett 2013, Brennan et al. 2006, Martine et al. 2011, a putatively natural group including the cultivated eggplant (Solanum melongena L.) and a number of other non-Australian species. Nine of the morphologically androdioecious -but functionally dioecious -spiny solanums known in Australia at that time were included by Symon in this group, as were a set of species that exhibit andromonoecy, with each infl orescence consisting of a basal hermaphroditic fl ower accompanied by numerous staminate fl owers that rise above it in a distal cyme.
Later work employing molecular phylogenetics , 2009, Vorontsova et al. 2013 showed that section Melongena sensu Symon (1981) was polyphyletic. A set of Australian andromonoecious species, however (including S. chippendalei Symon, S. diversifl orum F. Muell. and S. beaugleholei Symon), represent a moderately supported grouping identifi ed by Martine et al. (2006) as the "bush tomato clade" -a fi nding that supports inferences by Symon (1981), Whalen (1984) and Bean (2004) regarding their similarities and relatedness. One species inferred to be in this clade, S. eburneum Symon, was among the many new species described by Symon (1971) during his long career.
Apparently endemic to a small region of the Northern Territory around the East Baines River, S. eburneum appeared to Symon to be restricted to gray clay soils. Meanwhile, botanists collecting in surrounding areas, including Pete Latz (NT Herbarium, Alice Springs), had observed populations across a broader range that were recognizably distinct from the similar S. eburneum. A specimen sent by Latz to Symon in 1976 led the latter botanist to conclude that it matched "no very satisfactory name" used at the time and that he could "do no better than affi n. eburneum" (excerpt from letter on specimen NTD9293, P.K. Latz 5325). Field identifi cation was also sometimes challenging, especially along the Bullita Stock Route where the range of the apparently more widespread morphotype comes into contact with S. eburneum in Judbarra/Gregory National Park. Still, the morphotype was recognizable enough to be identifi ed for years on collection labels as S. sp. "Bullita" (Latz 12401) in reference to Latz's assignment of that identifi er to the putative new taxon beginning in the early 1970s. Until now, however, no morphometric comparison between the two morphotypes has been undertaken.
In 2014, CTM and family surveyed the vicinity of the East Baines/Victoria Highway/Bullita Stock Route region for occurrences of S. eburneum and S. sp. "Bullita." Seeds of typical S. eburneum (from the type locality as per Symon 1981) and S. sp. "Bullita" populations were collected and then grown at Bucknell University, where morphometric analyses were conducted on the two taxa in order to defi ne the diff erences between them and to determine the novelty of S. sp. "Bullita." We here describe this species as new, and contrast both its morphology and distribution with its close relative S. eburneum.

Morphological comparisons
In early May 2014, populations of S. eburneum were visited along the Victoria Highway corridor between Timber Creek and the Northern Territory border with Western Australia, including the type locality (Symon 1981) at the intersection of the highway and the East Baines River. Herbarium specimens and mature fruits were collected, with seeds extracted, dried, and stored for future use. Similar populations identifi ed as S. sp. "Bullita" were then collected in the same manner along the Bullita Stock Route in the Bullita section of Judbarra/Gregory National Park. Leaf material for each taxon was collected and dried in silica for subsequent DNA work (not included here).
To establish populations for morphometric analyses seeds were soaked for 24 hours in 1000-ppm gibberellic acid and sown in a controlled growth chamber environment at Bucknell University. Th ey germinated at a rate of 70-90% in 2-3 weeks, with seedlings transferred to individual pots in our research greenhouse after establishment. All plants were grown in a common soil mix under identical horticultural conditions. Th irty-eight vegetative and reproductive characters were measured across developmental stages in 41 cultivated plants. Th ese data were analyzed using ANOVA in the statistical software JMP-Pro 12 (SAS Institute Inc., Cary, North Carolina, USA) to defi ne characters diff ering between the taxa. Character diff erences were corroborated and "fi eld-truthed" by examination of herbarium specimens, including accessions held at the Northern Territory Herbarium, Palmerston (DNA) (See "specimens examined" and Appendix 1) from across the geographic range of each taxon ( Figure 1). Characterization of trichomes is based on descriptions by Bean (2012) for S. sp. "Bullita" derived from the author's examination of herbarium specimens.

Results
Th e ANOVA analyses found that of 38 discrete vegetative and reproductive characters that were measured, 17 diff ered with statistical signifi cance between Solanum eburneum and S. watneyi (Table 1). An additional set of qualitative characters also proved useful for drawing distinctions between the taxa (Table 2). Diagnosis. Solanum watneyi is distinguished from S. eburneum by its weakly erect and sprawling habit, long internodes and fruiting pedicels, often scabrous dark green Table 1. Statistical comparison of characters, S. watneyi and S. eburneum. Single asterisk (*) indicates statistical signifi cance with a 95-99% confi dence interval; double asterisk (**) indicates statistical significance with >99% confi dence interval. SD = standard deviation. All measurements in cm, except trichome density (per 0.5 cm leaf disk) and number of seeds per fruit.  Description. Weakly erect to sprawling sub-shrub to 50-60 cm. Rhizomatous and apparently clonal. Stems slender, woody, often bending to ground, especially when weighted by fruits; initially single stemmed, with strong lateral branching beginning at ca. 5-10 cm; internode lengths on mature stems averaging about 4 cm. Overall plant aspect dark green to gray-green, with older stems eventually pea-green to yellow-brown; pubescent throughout with porrect stellate trichomes (stems, leaves, pedicels, calyces) with stalks 0-0.1 mm long, lateral rays 6-10, central ray (midpoint) 1-1.5 times as long as the lateral rays; pubescence of stems short and loose. Prickles sparse, straw-colored, straight, slightly widened at base, fi ne, 2-5 mm long, scattered on stems. Sympodial units difoliate, the leaves solitary or geminate. Mature leaves 12-16 cm × 2-6 cm, alternate, lanceolate-elliptic, with 6-8 pairs of primary veins; young leaves lighter green and gray-hairy but becoming dark green above, slightly paler beneath, both sides closely and densely stellate tomentose, older leaves becoming scabrous and uniformly dark on both sides, retaining dense tomentum along veins; base attenuate; margins entire, sinuate or occasionally shallowly 6-8 lobed; apex rounded, mucronate; scarcely armed along midvein beneath; petiole 3-3.5 cm long with few to no prickles. Infl orescence a supra-axillary andromonoecious cyme 9-11 cm long, consisting of a basal hermaphrodite fl ower and a distal group of 3-7 (usually) staminate fl owers (the most basal staminate fl owers occasionally expressing as hermaphrodite), typically 1-2 staminate fl owers open at a time, common peduncle typically 2-5 mm (-20) long, rachis slightly less tomentose than stems. Flowers 5-merous, heterostylous. Hermaphrodite fl ower ca. 2.5 cm below the staminate fl owers, opening fi rst; pedicel 2-3 cm long at anthesis, elongating further after fertilization, armed with prickles 2-5 mm long; calyx lobes 5-8 mm long, armed with long, straight prickles and stellate trichomes; corolla 4-5 cm in diameter, dusty purple, rotate; acumens ca. 2 mm; ovary glabrous, ca. 2 mm diameter at anthesis; style 11-12 mm (including capitate stigma), curved; stamens equal; fi laments ca. 1 mm; anthers 6 mm long, oblong-lanceolate to somewhat tapered, poricidal at the tips, in a loose anther cone. Staminate fl owers with pedicels 15-16 mm long, unarmed or with few prickles; calyx lobes 7-8 mm long with a few 3-4 mm weak prickles or prickles absent, the lobes ending with a slender fi liform acumen ca. 3 mm long, slightly refl exed; corolla 2.5-4.5 cm in diameter, dusty purple, broadly stellate to rotate; acumens ca. 0.5 mm; stamens of same proportions as in hermaphrodite fl ower; ovary, style, and stigma vestigial and not exserted beyond the stamens. Fruit an ellipsoidal berry 2-2.5 cm long, 1.5-2.2 cm wide, light green with dark green stripes when young, maturing to lemon-yellow with faint brown stripes; fl esh fi rm, the locules 2, not liquid-fi lled and the internal cavity dry, the fruit wall ca. 0.5 cm thick, the fruits retained on plant after maturation. Fruiting pedicels 3.5-4.5 cm long, defl exed so fruits resting on ground at maturity. Fruiting calyx enclosing and exceeding fruit in early development, eventually covering up to 2/3 of developed fruit, the lobes long-acuminate, blunt-tipped, and weakly refl exing as fruit matures, short stellate-pubescent and armed with sharp spines 2-5 mm long, these single or paired along the calyx sutures, the lobes often fused in a 3+2 or 4+1 arrangement. Mature seeds up to ~100 per fruit, 2.5-3.5 mm, tan to dark brown, fi nely reticulate.

S. eburneum S. watneyi
Distribution and ecology. Solanum watneyi is presently known from a small range of localities in the sub-arid, monsoon-infl uenced, zone of northwestern Northern Territory (Fig. 1) at elevations around 100-150 m. Th e species is locally abundant along the Bullita Stock Route and nearby areas, but abundance elsewhere is not well known. Solanum watneyi appears to be associated, at least in Judbarra, with the "Ridges, Hogbacks, Cuestas, and Structural Plateaux" land unit defi ned by Brocklehurst et al. (1996), where it occurs in open Eucalyptus woodlands. Field collections have been made in well-drained limestone-based soils variously described as being sandy, sandyloamy, clayey-loamy, and loamy. Collections from 70-80 km east of the Bullita section suggest that the species range extends eastward into the Wanimyn Trust and the landscapes around the Buchanan Highway; and collections from Limbunyah Station hint toward a westward range extending toward the Western Australia border (Fig. 1). Th e areas where S. watneyi has been collected most frequently are along graded roads (Fig. 2D), suggesting that this taxon is disturbance-adapted. Although collections have been made in areas that experience bushfi res, specifi c fi re ecology adaptations are unknown. Th e pollination biology has not been studied, but the fl owers are likely buzz pollinated (see Anderson and Symon 1988) -as evidenced by anthers with terminal pores and the need for physical manipulation of anthers during hand pollinations in cultivation. Seed dispersal appears to be biotic given the fl eshy nature of the fruits, which ripen to yellow at maturity. Symon (1979) included S. eburneum in a group of solanums with fi rm yellow berries likely to be dispersed by mammals or larger birds; S. watneyi fi ts this same profi le.
Phenology. Most fl owering specimens have been collected from March-May, at the transition from the wet to dry seasons, with fruiting specimens collected in April-June. Earlier fl owering collections (e.g. K. Brennan 9002, 27 Jan 2000) suggest that blooming may begin during the rainy season and extend into the dry season. Plants in cultivation began fl owering about 90 days after seed germination; and fruits took about 60 days to mature following hand pollination.
Etymology. Th e specifi c epithet of "watneyi" is inspired by the book and fi lm, Th e Martian, in which the protagonist fi nds himself stranded on Mars surrounded by the planet's harsh terrain and reddish soils. In a shelter, he manages to grow a crop of potatoes (Solanum tuberosum) before fi nally being rescued by his astronaut colleagues. We've chosen to name Solanum watneyi after this character, Mark Watney, in part because of the similarly reddish soils of its habitat and the congeneric nature of the potato -but, most notably, as a way to honor the creation of a sci-fi hero botanist by author Andy Weir (Weir 2013) and to acknowledge perhaps the fi nest paean to botanical science (and botanical fi eld work) that Hollywood has yet presented (see Martine 2015).
Preliminary conservation status. Based on IUCN Red List Categories (IUCN 2012), S. watneyi should be considered Data Defi cient (DD). While the species appears to be relatively widespread over a range of approximately 6000 km 2 , it has been collected in fewer than 20 localities. Th e small number of collections, coupled with the fact that populations often consist of multiple individuals, suggests that the species is common in some localities but uncommon on the regional and global scales (apparently of restricted distribution). Further data are required before a certain conservation status can be determined. Key populations are protected in Judbarra/Gregory National Park and appear secure.  Discussion. Solanum watneyi has been known for some time as a recognized morphotype, having been described by Latz as Solanum sp. "Bullita" in the early 1970s. Th is name has been used as an identifi er by other botanists since that time (e.g. Bean 2012), appearing on the labels of herbarium specimens that are similar to S. eburneum yet conspicuous enough in the characters identifi ed above to merit attention.
Previous studies inferred that Solanum eburneum is part of the "Bush tomato clade" (Martine et al. , 2009, a clade of andromonoecious species that appears to have arisen between 4 and 1.5 million years ago (Särkinen et al. 2013). Preliminary results from a large-scale next generation molecular study (Martine et al. in prep) infer that S. watneyi is most closely related to S. eburneum and S. chippendalei. Solanum watneyi stands out among this group based on morphological diff erences that include its conspicuously long fruiting pedicels and internodes, lighter purple corollas, and a sprawling habit that often fi nds mature fruits resting on ground.
In the western (Bullita) tract of Judbarra/Gregory National Park, S. watneyi is locally common along the roadsides of the Bullita Stock Route between the Victoria Highway and the old Bullita Homestead, where it has been collected numerous times. Its abundance along this dirt thoroughfare suggests that the species appears, like many solanums, to respond favorably to disturbance. In fact, it appears most vigorously on the edges of graded dirt roads, resprouting from rootstocks after passes from mechanized grading equipment (Martine, pers. obs.) While our ex situ morphometric analyses and most fi eld observations are consistent with the distinctive morphological gestalt of S. watneyi, collections from around the Bullita Station can be somewhat confusing -with vegetative characters approaching intermediacy between it and S. eburneum. A small set of ex situ crosses between them (Martine, unpublished) suggests that interspecifi c hybridization is possible. Symon (1981) described S. eburneum as a narrowly-occurring species of "broad, shallow, seasonally dry Melaleuca swamps or fl ats with heavy grey soils" around the East Baines River. Th is localized endemism aligns with the characteristic physiography described by Brocklehurst et al. (1996) for the East Baines corridor, with S. eburneum occurring on clays and lithosols found on alluvial plains and rises along the river course as it runs northwest out of the Bullita section of the park (and on Dick Creek to the west and Timber Creek to the east). Th e presence of S. watneyi on well-drained sandy-loamy soils on the (largely) southern edge of this range suggest that the two closely related taxa might have diverged on disparate soil types in the last few million years (dating based on Särkinen et al. 2013).
Recent land use history may have brought the species into secondary contact on the north-south edge of their ranges in the central area of the Bullita Section, where an underlying geology including sandstone, siltstone, and dolomite creates an especially complex physiography (Brocklehurst 1996). Th e remote Bullita cattle station, the homestead of which is now a park attraction and outstation site, operated in this area for several decades beginning in the early 1900s. Th e Bullita Stock Route, a graded access road running south from the Victoria Highway, closely parallels the East Baines River near the homestead -providing a corridor of frequently disturbed habitat by which S. eburneum and S. watneyi might move and come into contact with the aid of road grading equipment or, perhaps historically, livestock. We suggest that the combined eff ect of ancient-origin physiography and recent disturbance in this area has created a zone of hybridization that deserves further study.