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QUEENSLAND GOVERNMENT Department of Environment
QUEENSLAND GOVERNMENT
Department of Environment
A Journal of Plant Systematics
Queensland Herbarium
AUSTIRORAINER
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etsrbes.- 8:22. Sa Re ee od La a a a a Td Ch a eres eee PIMENTEL SES RR ENS F
Editorial Committee
L.W. Jessup (editor) R.J.F. Henderson (technical advisor)
B.K. Simon (technical advisor)
Word Processing
Suzanne MacArthur
Austrobaliey2 Vol. 1, No. 1 was published on 1 December 1977 Vol. 4, No. 4 was published on 20 February, 1997
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BAM naednsan speech,
Austrobaileya 5(1): 1-153 (1997) Contents
Application of the phylogenetic species concept: A botanical monographic perspective
ANT eS THEO 83 er eeetace. Cane ne Whe Gh 7 EM face ede alle MeOH. IA Oreo ETE PE a wel OA oath! Rees hata
A taxonomic revision of Dissiliaria F.Muell. ex Baill. (Euphorbiaceae)
Peel Wo BGPP uote otac'es ceed word cn 'y Sl eaves 4 awh bla dg eh teased als 4, batenig ellen
Three new species of Kontainea Heckel (Euphorbiaceae) from Australia and Papua New Guinea
PAGE Orley occ, cote loys oe eS Ae fas ote A re eke se de etapa mteunla, Seth tet den Mowe Ween ele hy
A revision of Rubus subg. Malachobatus (Focke) Focke and Rubus subg. Diemenicus A.R.Bean (Rosaceae) in Australia
PAO Ne EMCEE? oe cv cigs ce 40, bere het a4 va 8. eld eee eee NG EOP ACERT MARY (Cte ETE hod le ere eons
Madangia inflata (Asclepiadaceae: Marsdenieae), a new genus and species from Papua New Guinea
Paul I. Forster, David J. Liddle and Iris M. Liddle ................0......
Marsdenia jensenti (Asclepiadaceae: Marsdenieae), a new species from north-eastern Queensland
PAU SEOs ete ma splits betes ayeeclesk cee ahs Cason sialic 42)e atte dua GF. hd hhh ketene ul oe clerk
A conspectus of Astrotricha DC, (Araliaceae) in Queensland, including two new species
De GAT as. av cores then Oe Soria, are 4a RPE be ee ees eee Paechnek reel eta are ee
Cooktownia robertsii, a remarkable new genus and species of Orchidaceae from Australia
ACE hs RCT ae, Os cts ple, BPE ng Ae ative ARTEL ce anal top tcantitces £, butste eeey catten ee Ain ccind eh tice
A taxonomic revision of the genus Austrosteenisia R.Geesink (Fabaceae: Millettieae)
UB 8! 6 ee es eee ea, eee Se PRL. POR PP eae DEPP ey ee eee
Rothia indica subsp. australis A.E.Holland (Fabaceae: Crotalarieae), anew subspecies occuring in Australia
Fit gro SEG) fot) 5 ner tae Ts Re RL SP ee LP OP Py, ee a, RO Pe ee See
Notes on Caesalpinia subg. Mezoneuron (Leguminosae: Caesalpinioideae} in Australia
| Se28 ot ex | | oa) a er ee ee OE pS ea eae Oe PPM Pea UL ERP ees 20 eee a
Guioa sarcopterifructa (Sapindaceae): a new Australian species
Peter C. van Welzen ............0000cu eee Ah UPN Es oN ace Sp tgces atte ied
Capparis velutina (Capparaceae), a new species from south-eastern Queensland
Peralta hs be seal eee ee cen: ce ok Ne ee eee eee
(continued)
Austrobaileya 5(1): 1-153 (1997)
Notes on the naturalised Flora of Queensland, 3 Pauls POrstet ses gcip etsy tain audi go sd econ pes 4-3 Ga aac ee hd nbd hg ee AE OPK te tee Ss 113
A new combination in Dendrocnide (Urticaceae) in north Queensland dk. Jgckes and ht, “Plane ye 5 c tipacace es oo bs Sebas mre oe pats 4 ecb dobar ye aceeb bg aey 121
Notes on Eucalyptus ser. Psathyroxyla Blakely (Myrtaceae) and other ‘Ash group’ eucalypts PAINT EREADS. pocket ee eek esd Roe wiey catsatn tesa Sen peaks Baral nad ihichisuee tee a etaaad aged SES ely 4 125
Leptochloa southwoodii (Poaceae: Chloridoideae), a new species from south-east Queensland Neil Snow and Bryan K. Simon ....... cc ccc ccc ccc ee eee cece e teen eee ees , 137
Lilaeopsis brisbanica (Apiaceae), a new species from Queensland, Australia PAG ALISO 9) 8 Apcis Sromis ever be be sole ends Wohi eve Ts Ee eget Aine. gy ores treed ee Bin TR capt eee tn 145
Microcarpaea agonis (Scrophulariaceae), a new species from south-eastern Queensland PR SANT tee 8 Sas dep e AY Ee oe Rowe ne -& wine pe dS Beach ed eon 6 Se ae ta 4 hai ng a 149
BRR HEV IOUS Joke eps rtd, 5 8 hu gS aercess ae ee Sy Dl Le att ss Ms ik pb horth ledge a he ne ay SARS § enlace) hie Ae 153
“CASA Ee
Application of the phylogenetic species concept: A botanical monographic perspective.
Neil Snow
Summary
Snow, N. (1997). Application of the phylogenetic species concept: A botanical monographic perspective. Austrobaileya 5(1): 1-8. The diagnosis of phylogenetic species and infraspecific taxa is considered from the perspective of botanical monography. Diagnosing phylogenetic species using population aggregation analysis (PAA) cannot be done in the herbarium, as it is a population-based procedure. However, herbarium specimens can be aggregated into phylogenetic species based on the presence of fixed characters, and PAA methodology in the field is encouraged to the maximum extent possible. The importance of diagnosing phylogenetic species by ordinary morphological means is stressed. A species is thus an aggregation of sexual or asexual semaphoronts consistently diagnosable by a fixed character or combination of characters recognizable by ordinary morphological means. Application of this species definition will generally decrease species numbers in botanical monographs and could therefore overlook significant but non-fixed phenetic variation. Given the widespread existence of such variation it is concluded that the recognition of infraspecific taxa remains desirable. However, since varieties are explicitly subordinate to subspecies in the Code and imply hierarchical pattern where none is expected to exist, the recognition of only one infraspecific category is recommended.
Keywords: phylogenetics, phylogenetic species, species concepts, subspecies, varieties, botany, monographs,
Neil Snow, Washington University, Department of Biology, Campus Box 1137, St. Louis, Missouri 63130, U.S.A. and; Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. Current Address: Queensland Herbarium, Meiers Road, Indooroopilly Queensland 4068, Australia email: Neil Snow@env.qld.gov.au
We need good theoretical contexts in which to formulate operational interpretations in order to proceed with empiricism. — Estabrook (1972).
Introduction
Jn the current phylogenetic era the histories of biological lineages have become of paramount importance for interpreting biological pattern and process. The impact of phylogenetic thinking has necessitated a critical re-evaluation of every important concept in systematics (Davis 1995; Harvey et al. 1996; Snow 1996).
For example, the phylogenetic perspective has spawned the concept of phylogenetic species (Nelson & Platnick 1981; Cracrait 1983, 1989; Donoghue 1985), the definition and identification of which have been controversial (Nixon & Wheeler 1990;
Accepted for publication 20 June 1997
de Queiroz & Donoghue 1990b; Wheeler & Nixon 1990). One debate has surrounded ‘history’ versus ‘character’ based approaches for diagnosing phylogenetic species (Mishler & Brandon 1987; de Queiroz & Donoghue 1988; Nelson 1989; Nixon & Wheeler 1990; de Queiroz & Donoghue 1990a,b; Baum & Donoghue 1995; Luckow 1995). A second debate has been whether phylogenetic species must be monophyletic (see Crisp & Chandler 1996). Another recent idea originating from the phylogenetic perspective is a methodological approach by which phylogenetic species can be recognized (Davis & Manos 1991; Davis & Nixon 1992). Termed ‘population aggregation analysis’ (henceforth PAA) and formulated largely in the context of isozyme data, PAA
2
tabulates populations for the presence or absence of characters and then iteratively ageregates these populations based on whether characters have gone to fixation (Davis & Nixon 1992; Davis & Goldman 1993).
McDade (1995) recently provided a detailed review of the monographic literature regarding species concepts and their application in practice (see also Hamilton & Reichard 1992). Although other botanists have considered the species problem from the phylogenetic perspective (Crisp & Weston 1993; Crisp & Chandler 1996), only two monographs cited by McDade (1995) (Soreng 1991: 508; Luckow 1993: 38) explicitly used a phylogenetic species concept sensu Nixon and Wheeler (1990). To the best of my knowledge, no papers published after the period covered by McDade (1995) addressed either the application of population aggregation analysis (Davis & Nixon 1992) using morphological characters, or the recognition of phylogenetic species in the herbarium. In addition, with the exception of Soreng’s (1991) detailed rationale, the issue of recognizing infraspecific taxa under a phylogenetic species concept largely has been overlooked (Luckow 1995).
Having recently completed a phylogenetically-based monograph of the grass senus Leptochloa P.Beauv. (Snow 1997) using standard approaches (herbarium, field, ereenhouse), I had to consider two issues relevant to the botanical monographer: 1) how to diagnose phylogenetic species relying mostly on herbarium specimens using morphological characters, and 2) the theoretical and pragmatic considerations of whether to recognize infraspecific taxa. [ here outline the rationale I followed, and which others may wish to follow, in the hope of stimulating further discussion on these topics.
Phylogenetic species and their recognition in botanical monographs
Under a phylogenetic perspective a monographer may wish to use a phylogenetic species concept (hereafter “‘PSC’) (Cracraft 1983, 1989; Donoghue 1985; Nixon & Wheeler 1990). A phylogenetic species has been defined
Austrobaileya 5(1): 1-8 (1997)
as ‘the smallest aggregation of populations (sexual) or lineages (asexual) diagnosable by a unique combination of character states im comparable individuals (semaphoronts)’ (Nixon & Wheeler 1990). Contrary to some views (de Queiroz & Donoghue 1988; Vrana & Wheeler 1992), it has been argued that the terms monophyly and paraphyly are not applicable at and below the species level (Nixon & Wheeler 1990; Davis & Nixon 1992). This follows the reasoning of Hennig (1966) that only entities expected to have hierarchical relationships, and not tokogenetic relationships, should be included in cladistic analyses.
An influential paper of Du Rietz (1930: 347) foreshadowed the need for population- based sampling as a necessary empirical antecedent of forming species boundaries. Population aggregation analysis (Davis & Nixon 1992) is not the first time that fixation of characters has been deemed necessary for the recognition of taxa (Davis & Heywood 1973: 78), nor is it the first methodological procedure linked directly to a particular species concept. As the best known example, gene flow and reproductive isolation are linked to the biological species concept, but these criteria are now rarely used in practice to diagnose species boundaries and are thus essentially non-operational (Whittemore 1993; Brookes 1996). In contrast, PAA provides precise methodological criteria by which phylogenetic species can be diagnosed, and in many cases it will be highly operational (but see below). Within the assumptions of the phylogenetic paradigm the logic and methodology of PAA is internally consistent. In many cases, however, the herbarium systematist will be able to apply PAA only under certain constraints.
One constraint of applying population ap oregation analysis 1s that, with the exception of small genera that occur 1n relatively close proximity to the monographer, PAA will often be logistically infeasible. This is particularly true for species-rich genera and those with intercontinental distributions. Few researchers have adequate time or funding for the necessary travel and intensive population sampling mandated by a strict adherence to PAA methodology. These realities acknowledged,
Snow, Phylogenetic species concept
monographers probably should collect using PAA methodology to the maximum extent possible; that is, sample multiple genotypes for a given population. This is particularly true for problematic species that appear to lack fixed characters or combinations of characters. The data collected from multiple genotypes in a population (hence, independent samples) are systematically informative because they permit analysis of intrapopulational variation, even if its evaluation is non-statistical. Such data are a critical consideration under the phylogenetic species concept, since fixed characters (100% occurrence) at the population level provide the basis by which phylogenetic species are recognized (Crisp & Chandler, 1996). For the herbarium taxonomist, however, the analysis of intrapopulational variation generally has been possible only when species were known from a single locality (e.g., tropical genera such as Cryptanthus Otto & A. Dietr. or Anthurium Schott.; see Appendix 1).
The second constraint regarding the recognition of phylogenetic species bears on what can happen when the PSC is pushed to its reductionist or aggregationist limits. The use of fixed characters to recognize phylogenetic Species may increase (Cracraft 1992) or decrease (Davis & Nixon 1992) the total number of species recognized. As previously argued (e.g., Olmstead 1995), a species could be recognized at the reductionist extreme on the basis of a single fixed nucleotide base pair. However, such a species would not be recognizable by ordinary means. A potential problem at the aggregationist extreme could be the lumping of multiple but phenctically distinct entities due to the absence of any fixed character(s) (see below). Those working with large tropical groups will be keenly aware of this potential problem. An aggregationist extreme also could overlook at least two important considerations in plant biology: 1} the existence of morphological outliers resulting from developmental abnormalities, which are not infrequent in plants, and 2) intermediates due to interspecific hybridization. However, biological reality does not reside within agarose gels or overly rigid theoretical constructs. Meaningful phylogenetic species, to the herbarium systematist, the ecologist,
3
and the community at large, must be those diagnosable by ‘ordinary means’ (Cronquist 1978). Thus, the second constraint of PAA methodology is that the monographer must recognize phylogenetic species using ordinary morphological means. To the botanical monographer, a phylogenetic species is an ageregation of sexual or asexual semaphoronts consistently diagnosable by a fixed character or combination of characters recognizable by ordinary morphological means. Users of herbarium-based monographs must realize, however, that additional but cryptic lineages may exist within morphologically based species. In Leptochloa an example would be the tetraploid, hexaploid, and octoploid levels occurring in the neotropical species Leptochloa dubia (Nees) Kunth (Gould 1975; Snow 1997).
Given the inability to always adhere strictly to the methodology of PAA, yet acknowledging the decided preference for phylogenetic species diagnosable by ordinary means, the best option for the botanical monographer to approximate the recognition of phylogenetic species in the herbarium is to search for a fixed character or.suite of characters by which a species can be consistently recognized. The emphasis on fixed (versus polymorphic) characters derives from the fact that only fixed characters are capable ofreliably suggesting that the relationships between taxa bearing them will be hierarchical, a necessary assumption of cladistic methodology (Hennig 1966; Davis & Nixon 1992; Doyle 1995). Fixed characters need not be qualitative; quantitative characters showing consistent gaps (and not merely differences in mean values) are also admissible diagnostic features (Thiele 1993; Luckow 1995). The monographer can thus aggregate herbarium specimens into phylogenetic species based on fixed characters observable by ordinary means. This approach is nothing new in monography, other than perhaps the requirement of character fixation. Differences of opinion regarding what constitutes ‘ordinary means’ will surely exist, but I see no reason to exclude characters that require up to 30x magnification, which is readily available in most herbaria.The ageregation of specimens based on character fixation is a repeatable and testable approach
4
(Snow 1996) that allows future workers to test hypotheses of species circumscription. With this approach a phylogenetic species is what the data (1.e., the characters) warrant, not merely what a competent systematist says it is.
Using the criteria for recognition of phylogenetic species outlined above, my revision of Leptochioa resulted in a decrease in species number (Snow 1997). Application of the PSC using field and herbarium data required merging the Australian taxa formerly known as Leptochloa ciliolata (Jedw.) S.T.Blake, L. decipiens (R.Br.) Stapf ex Maiden, and L. peacockii (Maiden & Betche) Domin (Lazarides 1980; Stanley & Ross 1989; Wheeler et al. 1990; Simon 1993) into a single species (Snow 1997). The necessity to lump occurred despite the fact that three readily distinguishable entities can exist in considerable numbers sympatrically within a few metres and maintain attributes which, in that locality, make them diagnosably distinct (Snow pers. obsv.). However, the attributes by which these taxa have been recognized can also break down in the field (Snow pers. obs.), and significant numbers of intermediates exist as herbarium specimens. Thus, whereas populations can be locally distinct, they may not be globally distinct. Since locally distinct populations not globally fixed for a character or combination of characters still probably reflect underlying genetic variation, it becomes necessary to consider whether infraspecfic taxa should be recognized.
The recognition of infraspecific phylogenetic taxa in botanical monographs
Theoretical considerations of infraspecific taxa have been largely overlooked during discussions of species concepts rooted in the phylogenetic paradigm (Luckow 1995). Many (Stebbins 1950; Wilson & Brown 1953; Grant 1981; Mayr 1982) but not all (Du Rietz 1930; Raven 1962; Raven & Raven 1976) pre-cladistic discussions of infraspecific taxa were linked to the ‘biological’ species concept, or BSC. Reproductive isolation and gene flow, the central tenets of the BSC (Whittemore 1993), cannot be invoked under the phylogenetic paradigm, given its different
Austrobaileya 5(1): I-8 (1997)
assumptions (Luckow 1995). If they are to be recognized, phylogenetic subspecies or varieties, like their ‘biological’ counterparts, probably will lack consistent criteria for their recognition (Darwin 1859; Grant 1981; but see Pimentel 1959).
Although cladistic approaches are routinely used by population geneticists below the species level to trace gene genealogies (Avise 1989; Crandall & Templeton 1993; Doyle 1995; Maddison 1995; Moritz & Hillis 1996), for the botanical monograph and the phylogenetic analyses on which the supraspecific groups will be based, phylogenetic species are the minimal units of cladistic analysis. However, that does not necessarily make phylogenetic species the minimal taxa. It is possible to formally recognize phenetic taxa below the species level which, unlike phylogenetic species, are not expected to be hierarchically related. This seems particularly defensible if the monographer has field experience with the group, since variation easily detected in the field is frequently not captured on pressed herbarium specimens. For example, in the orchid genus Cryptocentrum Benth., leaf phyllotaxis (spiral/distichous), mnfloresence position (erect/pendant), and floral bracts (spathaceous/tubular) are diagnostic and fixed characters readily observable in the field, yet are cryptic on herbarium sheets (G. Carnevali pers. comm.).
Returning to the earlier example of Leptochloa, application of the phylogenetic species necessitated lumping L. ciliolata, L. decipiens, and L. peacockii into one species. However, given that 1) others have recognized three phenetic entities (Simon 1993; Stanley & Ross 1990); 2) these phenetic entities can be diagnosably distinct sympatrically (Snow pers. obs.); but that 3) their distinctness can also break down in the field (Snow, pers. obs.), I recognized a single species with three subspecies (Snow 1997, 1n press). Infraspecific taxa can thus be phenetic groupings that presumably reflect genetically-based variation, but the patterns of which are non-fixed and non- hierarchical. Under this approach an infraspecific taxon will still be what a
1 YS ASE Ng ET
Snow, Phylogenetic species concept
competent systematist says itis (Hubbell 1954: 120; Hedberg 1957: 14; Raven 1962: 6), although one would hope that newly recognized. infraspecific taxa would still be supported with evidence (narrative comment or numerical data) of some sort (Hedberg 1957: 16). My view of infraspecific taxa as phenetic clusters that reflect underlying genetic variation is similar to the species concept outlined by Crisp and Weston (1993), except at a lower taxonomic level.
A question arises then regarding phenetic pattern: Should we formally recognize such pattern at the infraspecific level if 1t becomes badly blurred or non-diagnosable in a global context? In other words, should an infraspecific taxon be named if the taxonomic placement of some (or all) of its members requires knowledge of their geographical occurrence (du Rietz 1930; Stebbins 1950; Brown & Wilson 1954; Pimentel 1959; Mayr 1982)? For highly vagile organisms, such as the many r-selected annual species of Leptochloa, it 1s unreasonable to expect that infraspecific variants will always be restricted to a particular geographic domain (e.g., Snow & Simon in press). Characters, fixed or non-fixed, are heritable and emergent properties of populations or series of populations. Geographical distribution is not a heritable feature and does not necessarily reflect distinct genetic lineages, although it may reflect genetic variation. The actions of mutation, selection, drift, and lineage sorting collectively predict the existence of geographically localized variation in widespread species. If a putative infraspecific taxon cannot be distinguished independent of its geographical occurrence— that is, if no combination of non-fixed
characters exists by which a ‘specimen’ can be |
identified at the infraspecific level—then that taxon should not be recognized. Geographical distribution by itself is an invalid criterion for recognition of infraspecific taxa.
Finally, the monographer must consider how many infraspecific ranks should be recognised, The International Code of Botanical Nomenclature (Greuter et al., 1994) allows for names at two infraspecific ranks: subspecies, and at a subordinate level, varieties. However,
a
even one infraspecific level of non-fixed phenetic pattern implies fuzzy boundaries between the constituent infraspecific taxa. Under the phylogenetic species concept it is illogical to recognize more than one level of phenetic infraspecific pattern, since variety is subordinate to subspecies, and if both were recognized, it would imply hierarchical pattern where none was expected to exist (Nixon & Wheeler 1990). Complex patterns of variation can certainly occur (Gould & Johnston 1972; Thorpe 1983; Crisp & Chandler 1996), but those inclined to recognize both subspecies and. varieties under a phylogenetic species concept using ordinary means might consider whether they are attempting to recognize a degree of hierarchical pattern that does not really exist. As noted by others, the recognition of more than one infraspecific rank generally engenders confusion, especially to the non-specialist (Stebbins 1950: 32-33; Hedberg 1957: 15). With these considerations in mind it seems most appropriate to formally recognize ony one infraspecific category.
Conclusions
I agree with earlier views (Davis & Goldman 1993; Kellogg 1994; McDade 1995; Luckow 1995) that monographers should clearly outline their species concepts and methodological approaches so that future workers can improve upon extant taxonomies as additional data warrant. Otherwise, successive revisions are merely a chronological array of untestable opinions which, even if based on reliable data, are difficult to evaluate. The exposition presented here represents my own attempts to integrate and pragmatically apply recent theoretical advances in systematics to the writing ofa botanical monograph (Snow 1997),
Phylogenetic species are the minimal units amenable to cladistic analysis for the purposes of botanical monography in the phylogenetic perspective. A phylogenetic species is an aggregation of sexual or asexual semaphoronts consistently diagnosable by a fixed character or combination of characters recognizable by ordinary means. A single infraspecific level of non-fixed phenetic variation may be recognized taxonomically
6
provided the variation is not irretrievably obscured when the species is viewed globally. For the recognition of phylogenetic species fieldwork should focus on the population sampling of separate genotypes for poorly known species, and the populations evaluated using PAA to the maximum extent possible.
The methodology outlined here ts not meant to be critical of earlier work. Others will find much to disagree with regarding the logic and rationale outlined above for recognizing species and infraspecific taxa for a phylogenetically-based monograph. However, if this article stimulates further dialogue regarding these topics, particularly in the context of producing scientifically meaningful but useful botanical monographs, it will have served a useful purpose.
Acknowledgements
I thank the following for discussions regarding the ideas contained herein: J.Bradford, G.Carnevali, T.Croat, J. Davis, W.Hauk, P.Hoch, S.Malcomber, I[.Ramirez, and P.Raven. The comments of an anonymous reviewer were helpful. My research has been supported by the Missouri Botanical Garden (Andrew W.Mellon Foundation), a grant from the National Science Foundation to the Plant Biology Program at Washington University (St. Louis), and the National Geographic Society (NGS 5594-95).
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HEnnIiG, W. 1966. Phylogenetic Systematics. Urbana: University of Illinois Press.
Huse tt, T.H. 1954, The naming of geographically variant populations. Systematic Zoology 3; 113-121.
KEeLLoGG, E.A. 1994, What makes a good monograph? American Society of Plant Taxonomists Newsletter 8: 12-15.
LAZARIDES, M. 1980. The genus Leptochioa Beauv. (Poaceae, Eragrostideae) in Australia and Papua New Guinea. Brunonia 3:247-269,
Lucxow, M. 1993. Monograph of Desmanthus (Leguminosae-Mimosoideae), Systematic Botany Monographs Vol. 38.
7
— 1995. Species concepts: assumptions, methods, and applications, Systematic Botany 20: 589-605.
Mapopison, W.P. 1995. Phylogenetic histories within and among species. Pp. 273-287 in P. C. Hoch & A. G. Stephenson (eds.), Experimental and Molecular Approaches to Plant Biosystematics. Monogr. Syst. Bot. Missouri Bot. Gard. 53.
Mayr, E. 1982. The Growth of Biological Thought: Divergence, Evolution, and Inheritance. Cambridge: The Belknap Press of Harvard University.
McDapg, L.A. 1995. Species concepts and problems in practice: insight from botanical monographs. Systematic Botany 20: 606-622.
Misuuer, B.D., & Brandon, R.N. 1987. Individuality, pluralism, and the phylogenetic species concept. Biology & Philosophy 2: 397-414.
Moritz, C., & Hitus, D.M. 1996, Molecular systematics: context and controversies. Pp. 1-13, in D.M.Hillis, C.Moritz, G.K.Mable (eds.), Molecular Systemtatics, Second Edition. Sunderland, Massachusetts: Sinauer Associates, Inc.
Netson, G. 1989. Cladistics and evolutionary models, Cladistics 3: 275-289.
NELSON, G., & PLATNICK, N. 1981. Systematics and Biogeography: Cladistics and Biogeography. New York: Columbia University Press.
Nixon, K.C., & WHEELER, Q.D. 1990. An amplification of the phylogenetic species concept. Cladistics 6:211-223.
OLMSTEAD, R.G. 1995. Species concepts and plesiomorphic species. Sysetmatic Botany 20: 623-630,
PIMENTEL, R.A. 1959, Mendelian infraspecific divergence levels and their analysis. Systematic Zoology 8: 139-159.
RAVEN, P.H. 1962. The systematics of Oenothera subgenus Chylismia. University of California Publications in Botany, Vol. 34: 1-222.
RAVEN, P.H., & Raven, T.E. 1976. The genus Epi/obiun in Australasia. New Zealand Department of Scientific and Industrial Research, Bulletin 216.
SIMON, B.K. 1993. A key to Australian Grasses. Second Edition. Brisbane: Queensland Department of Primary Industries.
Snow, N. 1996. The phylogenetic paradigm of comparative biology — a response to Hedberg. Taxon 45: 85-90.
8
—— 1997, Phylogeny and Systematics of Leptochloa P. Beauv. sensu lato (Poaceae, Chloridoideae). Ph.D. Dissertation, Washington University, St. Louis.
—— In press. Nomenclatural changes in Leptechloa P. Beauv. sensu lato (Poaceae, Chloridoideae).
(Novon)
— Simon, B.K. In press. Australian distribution of the weedy neotropical grass Leptochloa fusca subsp. uninervia, with an updated key to Australian Leptochloa (Poaceae, Chloridolideae). (Austrobaileya)
SORENG, R.J. 1991. Systematics of the ‘Epiles’ group of Poa (Poaceae). Systematic Botany 16: 507-528.
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STEBBINS, G.L. 1950. Variation and Evolution in Plants. New York: Columbia University Press.
Appendix 1. The sampling of numerous genotypes from multiple populations could quickly result in curatorial problems for a given institution. To alleviate curatorial congestion, I suggest a separate collection number be assigned to each genotype and that specimen labels clearly reflect the genotypic uniqueness of each specimen. For example, a numbering scheme for a given population might be Collector et al. 7249-A, Collector et al. 7249-B, etc., with a statement on the label
Austrobaileya 5(1): 1-8 (1997)
THIELE, K. 1993, The holy grail of the perfect character: the cladistic treatment of morphometric data. Cladistics 9: 275-304.
THorpg, R.S. 1983. A review of the numerical methods for recognizing and analyzing racial differentiation. Pp. 404-423 in Numerical Taxonomy, (J. Felsenstein, ed.). Berlin: Springer-Verlag.
VRANA, P., & WHEELER, W. 1992. Individual organisms as terminal entities: laying the species problem to rest. Cladistics 8: 67—72.
Wueecer, D.J.B., Jacoss, S.W.L., & Norton, B.E. 1990. Grasses of New South Wales. Second Edition. Armidale: The University of New England.
WHEELER, Q.D., & Nixon, K.L. 1990. Another way of looking at the species problem: A reply to de Queiroz and Donoghue. Cladistics 6: 77-81.
Wuirremore, A.T. 1993. Species concepts: a reply to Ernst Mayr. Taxon 42: 573-583.
Witson, E.O., & Brown, W.L. Jr. 1953. The subspecies concept and its taxonomic application. Systematic Zoology 2: 97-111.
reading “Each letter designates a different genotype’. Upon completion of the monograph n-1 genotypes from each population could be distributed to other herbaria following standard procedures. An indication on the labels of the herbaria to which duplicates will be distributed would assist future workers. Given its relative ease collectors should also consider collecting fresh leaf matertal suitable for molecular studies in silica gel.
A taxonomic revision of Dissiliaria F.Muell. ex Baill. (Euphorbiaceae)
Paul lL. Forster
Summary
Forster, Paul I. A taxonomic revision of Dissiliaria F Muell. ex Baill. Euphorbiaceae), Austrobaileya 5(1): 9-27 (1997). The genus DissiliariaF Muell. ex Baill. is revised. Dissiliariais endemic to Australia with six species from eastern Queensland, D. baloghioides F.Muell. ex Baill., D. indistincta P.I.Forst. sp. nov., D. laxinervis Airy Shaw, D. muelleri Baill., D. surculosa P.1.Forst. sp. nov. and D. tuckeri P.IJ.Forst. sp.nov. All species are described and illustrated, with notes on distribution, habitat, typification and conservation status. The names D. baloghioides and D. muelleri are lectotypified.
Keywords: Euphorbiaceae, Dissiliaria - Australia; Dissiliaria baloghioides, Dissiliaria indistincta, Dissiliaria laxinervis, Dissiliaria muelleri, Dissiliaria surculosa, Dissiliaria tucker.
Paul I. Forster, Queensland Herbarium, Meiers Road, Indooroopilly, Queensland 4068, Australia
Introduction
The genus Dissiliaria was described by Baillon (1867) who recognised two species D. baloghioides F.Muell. ex Baill. and D. muelleri Baill. Baillon provided descriptions and cited types for the two species he named, but did not provide a formal generic description of his genus although he did extensively discuss its affinities and morphological characters. Airy Shaw (1981) considered that Baillon’s account did not constitute formal description; however, other authors (e.g. Bentham 1873; Chapman 1991; Webster 1994) and the compilers of
Index Kewensis all considered it to be valid. A —
third species, D. laxinervis, was described by Airy Shaw (1980).
Dissiliaria was included both by Webster (1994) and Levin & Simpson (1994) in Euphorbiaceae subfamily Oldfieldioideae Kohler & Webster, tribe Caletieae Muell.Arg., subtribe Dissiliariinae Pax & K.Hoffm. Other genera included in subtribe Dissiliariinae are Austrobuxus Mig., Canaca Guillaumin (included with a query inAustrobuxus by Levin & Simpson (1994)), Choriceras Baill., Longetia Baill. ex Muell.Arg., SankowskyaP.I.Forst. and Whyanbeelia Airy Shaw & B.Hyland (Levin & Simpson 1994; Webster 1994; Forster 1995). A comparison of macromorphological
Accepted for publication 16 May 1997
characters for genera in the Dissiliariinae, together with a generic key may be found in Forster (1995). The genus Dissiliaria 1s characterised by the plants being dioecious and stipulate; the male flowers having 3+3 sepals, a hairy receptacle, glandular disk absent, 8—26 stamens, anthers oblong, pollen spiny and pistillode absent; the female flowers having 3+3 sepals, a glandular disk present and the styles linear in outline; the fruit globose and having laterally compressed seeds that have an entire caruncle. Anatomical features that distinguish species of Dissiliaria. from other taxa in the Dissiliariinae according to Hayden (1994) are “high rank venation, well-developed and somewhat oriented areoles, narrow ultimate tracheids in veinlets, fimbrial veins, wavy anticlinal walls of leaf epidermis cells, bundle sheath extensions, and libriform wood fibres”. Based on palynological, vegetative and anatomical characters, Levin & Simpson (1994: 226) produced a cladogram wherein Dissiliaria was basal in the Dissiliariinae, and more closely allied to Austrobuxus and Whyanbeelia than to Longetia and Choriceras.
All the species of Dissiliaria are trees and grow in rainforest communities. Fertile specimens of Dissiliariahave proven extremely difficult to acquire as flowering and fruiting is irregular and some of the localities where the
10
plants occur are difficult to access when they are fertile. This situation was alluded to by Airy Shaw (1981). Nevertheless, there is now considerably more material to review than was available to Airy Shaw. Consequently it is possible to resolve the status of specimens originating from the populations in the Mcliwraith Range on Cape York Peninsula that Airy Shaw referred with doubt to D. muelleri. These are described as the new species D. surculosa. Airy Shaw also included a number of taxa within his concept of D. laxinervis. It is now possible to distinguish a further two new species allied toD. laxinervis in the strict sense with D. indistincta and D. tuckeri newly described in this paper. Hence, six species of Dissiliaria are recognised in the present account.
The species of Dissiliariareadily fall into two groups based on the morphological characters of leaf margin (entire or crenate) and fruits (strongly or weakly trilobate). Crenate leaves and strongly trilobate fruits occur in D. muelleri and D. surculosa P...Forst. Entire leaves and weakly trilobate fruits occur in the remaining species. The occurrence of crenate leaves is considered plesiomorphic and occurs in unrelated taxa in the Oldfieldioideae, although with a preponderance in the Dissiliarimae (Hayden 1994).
The different species of Dissiliaria occur in restricted and relictual populations with numerous disjunctions over some 15 degrees of latitude in eastern Queensiand. This indicates that the extant species are relatively ancient with distributions fragmented by unfavourable environmental conditions at some time in the past, resulting in their being restricted in present times to refugial niches. Patterns of dispersal are unstudied in detail, but all species exhibit septicidal dehiscence of the capsular fruits followed by ‘drop and roll’ dispersal of the laterally compressed seeds. It is common to see profuse seedling beds beneath large female trees indicating that little lateral movement of seed occurs. In some instances, populations may be found for considerable distances along creeks on alluvia, indicating secondary water transportation of seeds occurs.
Austrobaileya 5({1): 9-27 (1997) Materials and methods
This revision is based on herbartum holdings at BRI, CANB, CBG, NSW, MEL and QRS, photographs of material in BM, G-DC, K and P, and field collections and observations by the author. All specimens have been seen unless indicated n.v. Indumentum cover is described using the terminology of Hewson (1988), except that ‘scattered’ is preferred to ‘isolated’. Indumentum in Dissifiariauniformly comprises simple, multicellular trichomes, thus reinforcing the observations of Hayden (1994) and further refuting those of Rao & Raju (1985).
Conservation codings are allocated using the terminology as defined in the schedules of the Queensland Nature Conservation Act 1992. Rainforest classification follows the scheme proposed by Webb (1978). The ‘Wet Tropics’ is defined as the area of north-eastern Queensland that encompasses the ‘hot, humid vine forests’ from near Cooktown in the north to Paluma in the south (Webb & Tracey 1981; Barlow & Hyland 1988).
Taxonomy
Dissiliaria F.Muell. ex Baill., Adansonia 7: 366 (1867). Type: D. baloghioides F.Muell. ex Baill. lecto, fide Webster (1994: 58).
Derivation of name: trom the Latin dissilio (to fly apart) and - aria (resembling or provided with), alluding to the dehiscence of the fruit capsule.
Trees, evergreen, perennial, dioecious; stems and foliage without latex. Indumentum. of simple, multicellular trichomes; glandular trichomes absent; stinging trichomes absent. Stipuies entire, inconspicuous or conspicuous, deciduous. Leaves opposite, petiolate, elobate, penninerved; margins entire or crenate. Inflorescences axillary, racemose, solitary or fascicled, unisexual with flowers in bracteate clusters. Male flowers pedicellate; sepals imbricate, 3+3; petals absent; receptacle convex; disk absent; stamens 8—26; filaments free, filiform to flattened-filiform, attached to slightly convex receptacle; anthers dorsifixed, bilobate; thecae oblong and longitudinally
Forster, Dissiliaria
dehiscent; pistillode absent. Female flowers pedicellate; sepals imbricate, 3+3; petals absent; disk shortly annular or cupular; ovary 2 - or 3-locular; locules biovulate; styles 3, erect, linear-subulate, entire, shortly connate at base.
il
Fruit capsular, trilobate, with surface smooth to muriculate, dehiscing septicidally into 3 bivalved cocci. Seeds semi-elliptic in outline, laterally compressed; testa crustaceous; albumen fleshy; caruncle entire, non-arilloid; cotyledons broad, flat.
Six species endemic in Australia.
Key to species of Dissiliaria
1. Leaves chartaceous, < 9.5 cm long, crenate
Leaves coriaceous, 9.5—18 cm long, entire
2. Leaf lamina narrowly elliptic with 17—23 marginal teeth per side of the midrib; typ acuminate; male flower sepals obovate, 0.8—-1.2 mm
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Gee en aS ates eases age 6. D. surcuiosa
Leaf lamina broadly ovate to rhombic-ovate with 14-16 marginal teeth per side of the midrib; tip acute, obtuse or retuse; male flower sepals
broadly elliptic, 1.4-1.5 mm wide.......
ese So Bee RA Sy Ey 5S. D. muelleri
3, Stipules lanceolate-obovate; stipular scar 1.8~2.2 mm wide .............. 4. D. tuckeri Stipules linear or deltoid-lanceolate; stipular scar 0.5—0.8 mm
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4, Stipules linear, c. 4 mm long; interlateral venation prominent below with ali connecting veins raised; pedicels 2.5-9 mm long .............. 1. D. baloghioides
Stipules deltoid-lanceolate, 1—-1.5 mm long; interlateral venation weakly
visible below with only the outer connecting veins raised; female pedicels
10-40 mm long ..... 0... 00 eee eee
5. New foliage green to pale green-pink; styles c. 6 mm long, erect,
New foliage bright pink-brown; styles 2.5-4mm long, strongly recurved, connate for c. | mm; stamens 16-18; anthers 0.8-1.2 mm
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1. Dissiliaria baloghioides F.Muell. ex Baill., Adansonia 7: 366 (1867). Type: Queensland. Moreton District: [label in Leichhardt’s hand] “scrub towards the Bunyas, 18/9/1843”; [printed label] Moreton Bay; [label in Leichhardt’s hand] “Wood 14....the blackfellows make their boomerangs of it”; [label in unknown hand} “n.328h. Leichhardt Moreton Bay” (ecto [here designated]: P n.v., photo at BRI).
Iflustrations: Francis (1981: 131, 132); Hauser (1992: 216).
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3. D. indistincta
Tree to 36 m high. Trunk slightly flanged at base. Bark rough, flaky-scaly in patches, grey to grey-brown; blaze pink. Branchlets + rounded, glabrous or with only scattered trichomes when young, lenticellate with age. Stipules linear, c. 4 mm long and 0.7 mm wide, with scattered trichomes. Leaves glabrous; new fohage pale pink; petioles 4-10 mm long, 0.8—1 mm wide, glabrous; laminas ovate, obovate or elliptic, 50-120 mm long, 15-70 mm wide, coriaceous, with margins entire and venation composed of 8 or 9 lateral veins per side of midrib and interlateral reticulate tertiary veins; upper surface shining
12
dark green with venation + obscure; below glossy pale green with venation prominent with all interlaterals raised; tip acute to obtuse; base attenuate, obtuse or truncate. Male inflorescences in distal axils, up to 7 mm long; peduncles up to 1 mm long; bracts lanceolate, 1-1.5 mm long, 0.4-1.2 mm wide. Male flowers with pedicels 4-12 mm long, 0.5— 0.9 mm diameter, with sparse to dense golden trichomes; sepals elliptic, 1.8-4.5 mm long, 1.2-2 mm wide, with sparse trichomes; disk 2—2.5 mm diameter, with dense trichomes; stamens 15—20; filaments1.5—3.1 mm long; anthers 0.5—1 mm long, 0.3—-0.7 mm wide. Female inflorescences in distal axils, of solitary flowers with peduncles obsolete; bracts ovate, c. 2 mm long and 1 mm wide, with sparse trichomes. Female flowers with pedicels 2.5—9 mm long, 1.2—2 mm diameter, with dense golden trichomes; sepals elliptic to ovate, 3-4 mm long, 1.8-2.2 mm wide, with sparse to dense trichomes; ovary 2.5—3 mm diameter, with dense golden trichomes; styles 2.6—3.5 mm long, shortly connate for c. 1 mm in length, strongly recurved, papillose on upper surface, with dense trichomes on lower surface for entire length. Fruits globose, weakly trilobate, 12—21 mm long, 15—22 mm diameter, minutely muriculate; seeds 8-10 mm _ long, 4.5—-5.5 mm wide, 2—2.4 mm thick, smooth, shiny brown. Fig. 1.
Selected additional specimens examined: Queensland. NortH Kennepy District: Slaty Creek, near Proserpine, Michael 988 (BRI). SourH Kennepy Districr: Cherrytree Creek, Koumala, Nov 1987, Canning 305 (BRD); Foot of ranges W from Koumala, May 1921, Francis |AQ202261 | (BRI). Port Curtis District: Pine Creek, off lower reaches of Granite Creek, S.F. 391 Bulburin, 24°37’S, 151°33’E, Dec 1993, Forster 14580 et al, (BRD; S.F. 67, Bulburin, Sep 1983, Gibson 559 (BRI); Crossing Reserve R73 Miriam Vale, on Baffle Creek, 5 km E of Miriam Vale on road to Agnes Water, Jul 1978, McDonald 2347 & Stanton (BRI, CANB). Wins Bay District: Amamoor, May 1918, Forestry Dept. [AQ202271] (BRD; S.F. 50, 3.5 km S of Mt Urah, 25°52’S, 152°22’E, Nov 1988, Forster 4842 (BRI, CANB, MEL); S.F. 50 Glenbar, 1 km WSW of Mt Urah summit, 25°50’S, 152°20’E, Jan 1993, Forster 13126A & Machin (A, BRI, K, L, MEL, QRS), 13126B (BRI, K, L, MEL, QRS); Kin Kin, Eastern Branch road, 26°18’S, 152°53’°E, Feb 1995, Forster 16123A (BRI, MEL, NSW, QRS). Moreton District: Bahr’s Scrub, $ of Beenleigh, 27°43’S, 153°10’E, Jun 1983, Guymer 1857 (BRI, CANB, MEL); 7 km SE of Mt Nebo, Scrub Road, Brisbane Forest Park, 27°25’S, 152°50’E, Sep 1993, Halford Q1842 (BRI); Petrie Creek, West Woombye,
Austrobaileya 5(1): 9-27 (1997)
26°40’S, 152°57°E, Apr 1986, Sharpe 4324 & Guymer (BRI); Eumundi, May 1896, Simmonds 470 (BRD; Kelvin Grove, Sep 1920, White & Francis [AQ202263] (BRD; Highvale near Samford, 27°22’S, 152°49°E, Dec 1983, Williams 83091 (BRI).
Distribution and habitat: D. baloghioides occurs in the North Kennedy, South Kennedy, Port Curtis, Wide Bay and Moreton districts of Queensland in disjunct populations (Map 1). This species is more widely distributed than stated by Airy Shaw (1981), occurring from the Mt Dryander area in the north to near Brisbane in the south. Plants grow in notophyll vineforests on volcanic substrates derived from eranite or rhyolite and are often locally abundant, particularly on creek alluvia.
Notes: Baillon (1867) cited three elements in the protologue of D. baloghioides, namely “Leichhardt, 0.13, n.14 (1843), Moreton Bay” and “Fitzalan, Pine River, Port Denison’. [have not been able to locate the Fitza/an collection, but what appears to be part of the Leichhardt collections is extant at P. This P material has several labels attached, but has the necessary requirements of at least one of the cited numbers (n.14), the correct date (1843) and the correct locality (Moreton Bay). The Leichhardt collection (n.14) is thus selected as lectotype of the name.
Some collections of D. indistincta P.I.Forst. from Mt Dryander in the North Kennedy district have been misidentified in herbaria as.D. baloghioides; however this latter species is easily distinguished by the indistinct interlateral venation in the lower leaf lamina as well as several other features (Table 1). D. baloghioides does occur at Mt Dryander but psrows in drier communities on the western slopes of the mountain (M.C.Tucker, pers. comm. 1994), whereas D. indistincta is common along creeks to the south and south-east of its base.
Phenology: Flowers February to June; fruits February to December.
Conservation status: Widespread, but not common with several populations in conservation reserves (Forster et al. 1991). No conservation coding 1s thought necessary.
by CA Ts Te see er tee che Bee EAT AEE Pe CC Frae hE eC CRATE Sree a Pane A Maa! ed Meee
Forster, Dissiliaria [3
= * - a* Be", + 4 % * * = _ = ~;, + i = +s* “hg é wes” + f oer om . + + _ + ' = sl =. = * iat + «= i; s+ - a4 * * =. + et ¥ i + * fF - *. f =. * ~~ + me +, ~~“ _* * = 7 + * 7a, = * att . Tey 4" * ¥ é we ‘S oye,? * ae a = Fes 4 tat? La tite bit +, 73 7, Par or ty ¥, * + ? ¥ +4 + . av * z a #7 SF 1 - i ets 4 ¥ ¥ Fa * ¥ * ss i tf + ri \ a oe tt ¢ * * = Pat +
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US
eK
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ate # +a tt te me hat
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Fig. 1. Dissiliaria baloghioides. A. flowering twig x 1. B. detatl of adaxial leaf surface x 1. C. female inflorescence x 2. D, female flower x 6. E, male flower x 6. F. stamen x 8. G. lateral view of fruit x 1. H. apical view of fruit x 1.1. one bivalved coccus of fruit x 2. J. persisent columella of dehisced fruit x 2. K. lateral view of seed x 2. L. ventral view of seed x 2. A,B,E,F from Forster 13126B & Machin (BRD; C,D from Forster 13126A & Machin (BRI); G,H from Smith {AQ316164] (BRD; E-L from Williams 83091 (BRI). Del. W. Smith.
14
Ethnobotanical use: D. baloghioides was reputedly used by aboriginal people in the Bunya Mountains area of south eastern Queensland to make boomerangs (label data with type). There are no extant populations of this species in the Bunya Mountains (cf. Forster et al. 1991) and it is likely that Leichhardt’s specimens came from further east, in the Blackall Ranges that were also once referred to as the Bunya Mountains (R.Fensham, pers. comm. 1997),
Etymology: The specific epithet refers to a superficial similarity to what is now known as Baloghia inophylla (G. Forst.) P.S.Green.
2. Dissiliaria laxinervis Airy Shaw, Muelleria 4; 220 (1980). Type: Queensland. Cook District: Claudie River, 12°50’S, 143° 20’E, 29 Jun 1972, B. Hyland RFK2578 (holo: K n.v.; iso: BRI, QRS).
Austrobaileya 5(1): 9-27 (1997)
Tree to 30 m high. Trunk buttressed at base, not suckering; bark rough and flaky in patches, blaze red. Branchlets + rounded, glabrous or with a few scattered trichomes when young, lenticellate with age. Stipules deltoid-lanceolate, 1—1.5 mm long, 0.5—0.6 mm wide, glabrous. Leaves glabrous; new foliage green or very slightly pale pink-green; petioles 7-9 mm long, c. 2 mm diameter, channelled on top, glabrous; lamina elliptic, 45-130 mm long, 30—70 mm wide, coriaceous, with margins entire and venation composed of 7 or 8 lateral veins per side of midrib and weakly developed interlateral reticulate tertiary veins; upper surface glossy dark green, with venation + obscure; lower surface pale green, with venation weakly developed with only the outer interlateral veins raised; trp acute to obcordate; base cuneate to rounded. Male inflorescences in distal axils, + sessile; bracts triangular, 1-1.2 mm long, c. 0.8 mm wide, glabrous; flowers in glomerules of 4-12. Male flowers with pedicels 22-30 mm long, c. 0.7 mm
Map I. Distribution of A Dissiliaria baloghioides, @ D. laxinervis and % D. tuckeri
Forster, Disstliaria
diameter, glabrous or with scattered hairs; sepals lanceolate-obovate, 4-5 mm long, 2.8—3 mm wide, glabrous or with scattered hairs; disk 3-4 mm diameter, with scattered to sparse short trichomes; stamens 22-26; filaments 1.2—1.8 mm long, c. 0.1 mm diameter, glabrous; anthers 1.3—1.7 mm long, 0.9-1 mm wide. Female inflorescences in distal axils, of solitary flower with peduncle obsolete; bracts lanceolate, c. 1.5 mm long and 0.4 mm wide,
15
glabrous. Female flowers with pedicels 17-40 mm long, c. 1 mm diameter, glabrous; sepals lanceolate, 2.8-4 mm long, 1.2—2.7 mm wide, glabrous; ovary globose, c. 2 mm long and 2 mm diameter, with dense yellow trichomes; styles c. 6 mm long, shortly connate for 2—2.5 mm at base, erect and with dense trichomes abaxially and shortly papillose apically. Fruits globose, weakly trilobate, 14-18 mm long, 15—18 mm diameter, minutely muriculate; seeds 8-10 mm long, 6.5—7 mm wide, 1.5—2.1 mm thick, smooth, shiny brown. Fig. 2.
Table 1. Comparison of morphological characteristics of entire-leaved species of Dissiliaria
Character
D. baloghioides D. laxinervis =D. tuckeri
D, indistincta
no. of lateral veins 8 or 9 7 ors $—10 S—/
per side of leaf lamina
midrib
stipule shape linear deltoid- lanceolate- deltoid- lanceolate obovate lanceolate
stipule size (mm) c,4 x 0,7 11.5 x 0.5-0.6 2.2-4 1.8-2.2 1-1.3 x 0.5-0.8
interlateral veins prominent indistinct indistinct indistinct
abaxially
stamen number 15-20 22—26 ? 16-18
style length (mm) 2.6—3.5 c. 6 44.5 2.94
styles recurved erect recurved recurved
styles connate atbase 1mm 2—2.5 mm ilmm 0.8—1 mm
female flower pedicel 2.5-—9 17-40 15-33 10-30
length (mm)
new foliage pale pink green orvery bright pale pink-brown bright red-pink green-pink
petiole dimensions (mm) 4-10 x 0.8-1 7-9 x ¢. 2 4-8 x] .5-2.5 3-7 x 1-2
16
Austrobaileya 5(1): 9-27 (1997)
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sot £7 ae af ie Pd hs = ont te a? r — aa $ He a ‘ = * ak | alt — = cat + os a 49.5 t "oe om, * ait + ores ai Dake ad ste? tianiie + ee, Py 725 we ant ix f btm dt A fF i, Pitan = . Paue? 4 aU Sissi
Vite, 3 te
c Zs Col a Lam
a aa 4 4 “1 i
Fig, 2. Dissiliaria laxinervis. A. twig x 0.4. B. male inflorescence x 1.5. C, male flower x 5, D. female flower = 5. FE, apical view of fruit x 1.5. F. lateral view of fruit x 1.5. G. ventral view of seed x 2.5. H. lateral view of seed x 2.5. A from Forster 13560 (BRI); B,C from Sankowsky 871 (QRS); D from Ayland 11516 (QRS); E-H from Gray 3626 (QRS). Del. W. Smith.
Forster, Dissiliaria .
Selected additional specimens examined: Queensland. Cook District: Iron Range, 12°46’S, 143°17’°E, May 1992, Cooper & Cooper 283 (QRS); Claudie River, 4 km ENE of Mt Tozer, Iron Range N.P., 12°44’S, 143°15’E, May 1992, Fell 2553 (BRI, QRS); Range 10 miles [16 km] NE of Iron Range, Apr 1944, Flecker N.Q.N.C.8689 (BRI, QRS); West Claudie River Scrub, 12°44’S, 143°14’E, Jul 1993, Forster 13560 et al. (BRI, MEL); Claudie River, 12°44’S, 143°15’E, Oct 1984, Gray 3615, 3626 (QRS); Claudie River, Jan 1982, Hyland 11516 (QRS); Claudie River, 12°44°S, 143°14’E, Sep 1988, Sankowsky 87] (QRS), 872 (QRS).
Distribution and habitat, D. laxinervis 1s known from a number of populations in the Claudie River catchment to the west of Iron Range on Cape York Peninsula (Map 1). Plants grow along permanent watercourses in mesophyll vineforest on alluvium derived from granite substrates.
Notes: D. laxinervis was described by Atry Shaw (1980) based on fruiting material. There is still a dearth of fertile herbarium material for this taxon, although itis now possible to provide descriptions of both male and female flowers here. Airy Shaw included material from the ‘Wet Tropics’ (1981) and Mt Dryander (annotations at BRI) within his concept of D. laxinervis, although he did comment that the collections from Gap Creek in the ‘Wet Tropics’ may be different. D. laxinervis is considered to occur only at Iron Range on Cape York Peninsula, and the southern populations mentioned by Airy Shaw are described in this paper asD. tuckeri and D. indistincta. They are compared with D. laxinervis in Table 1.
Phenology: Flowers have been collected in September and January; and fruits from May to October.
Conservation status: With the narrower circumscription ofD. laxinervis accepted here, the species is now confined to a single degree Square area on Cape York Peninsula. D. laxinervis 1s extremely common in the Claudie River Scrub and many plants are present in Iron Range National Park. No conservation coding 1s warranted.
Etymology: The specific epithet is dertved from the Latin /axis (ax) and nervis (nerve) and refers to the lax nervation in the leaf lamina of this species.
17
3. Dissiliaria indistincta P.I.Forst., sp. nov. affinis D. laxinervi Airy Shaw a qua frondescentia nova clara rosei-rubra, stylis brevioribus (2.5 — 4 mm longis) valde recurvatis et breviter connatis (per 0.8—I mm), et floribus masculinis staminibus paucioribus (16 —18) et antheris brevioribus (0.8—1.3 mm longis) differt. Typus: Queensland. Nortu KENNEDY District: Dryander Creek, left branch, base of Mt Dryander, 20°106’S, 148°34’E, 23 Feb 1994, PI. Forster 14856 & A.R. Bean (holo: BRI [3 sheets + spirit]; iso: A, B, BISH, CANB, DNA, G, Kk, L, MEL, MO, NSW, P, QRS, SING, US).
Tree to 25 m high or occasionally a shrub. Trunk often multistemmed, not butressed, not suckering; bark rough and flaky in patches; blaze pink. Branchlets + rounded, glabrous or with a few scattered trichomes when young, lenticellate with age. Stipules deltoid-lanceolate, 1—1.3 mm long, 0.5-0.8 mm wide, glabrous. Leaves glabrous, new foliage bright pink-brown; petioles 3—7 mm long, 1—2 mm wide, channelled on top, glabrous; lamina elliptic, oblanceolate, 30-130 mm long, 10-55 mm wide, coriaceous with margins entire and venation composed of 5-7 lateral veins per side of midrib and weakly developed interlateral reticulate tertiary veins where only the outer veins are raised; upper surface glossy dark green, venation + obscure; lower surface pale green, venation weakly developed; tip acute to obtuse; base cuneate to rounded. Male inflorescences in distal axils, + sessile; bracts lanceolate, c. 1.3 mm long and 0.8 mm wide, with scattered hairs; flowers in glomerules of 1~3, Male flowers with pedicels 9-21 mm long, c, 0.6 mm diameter, with scattered hairs; sepals broadly ovate to obovate, 2.8—3 mm long, 2—3 mm wide, glabrous; disk c. 2.5 mm diameter, with sparse to dense hairs; stamens 16-18; filaments 1.7—2.5 mm long, c. 0.2 diameter, glabrous; anthers 0.8—1.2 mm long, Q.8—1 mm wide. Female inflorescences in distal axils, of solitary flowers with peduncles obsolete; bracts lanceolate-triangular, 0.8—-5 mm long, 0.6-2 mm wide, glabrous. Female flowers with pedicels 10-30 mm long, 0.7-1.2 mm diameter, glabrous; sepals
18 Austrobaileya 5(1): 9-27 (1997)
CG EMELEM OY - * Jor v4 ie al Fe 7, a! at - 4A, # wks ae ee
IS
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oe ef ALA Lars
of ef
OS
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ae
Wt ry al a 1
in aif. “a ar 7 cs eee “eh ey le Me ry
— pel Sey mem Tem ey ie sag “te : at hs i iT “pati i — TNe eee be : ‘ih ' ’ A art " ] oe 1" Vipdneetes Fe RIO Ry it ere
i t : if | chiens me He Ht eT Cie ge pe BES +f pee : Bere EES einai Rey Fe “aS eons ye: Tikes 1 if a RR RR wee eee Ee t we Sa : ari = af : A ; . Z z [I 1 “< 4 , 3
z ee or ste Pati
+ : : ; z Shane 3 a AS toy oe aphedes a ara 7 7 cuts : : F a - yee ae Ay (5 the " 1 = : eae = SiMe aS l SY see Es te ena <Axseee al SES
Fig. 3, Dissiliaria indistincta. A. flowering twig x 0.5. B. detail of adaxial leaf surface x 1. C. maie flower x 5, D. stamen x 10. E. female inflorescence with two flowers x 2.5, F. lateral view of fruit x 2.5. G. apical view of fruit < 2.5. H. lateral view of seed x 10. I. ventral view of seed x 10. A, B, E from Forster 14856 & Bean (BRI); C, D from Forster 14857 & Bean (BRI), F, G from Forster 14302 (BRI); H, I from Sharpe 4175 (BRB. Del. W. Smith.
fagn ny SiS RASIURA SA SESS EEE AEE
ie Rivka ies enon ces nash ad cand gpg a aad
Forster, Dissiliaria
lanceolate to lanceolate - ovate, 1—S5 mm long, 0.5-4 mm wide, glabrous or with scattered hairs at base; ovary subglobose, 3—4.5 mm long, 3—5 mm diameter, with dense golden trichomes; disk annular to irregular cupular, 0.4—2 mm long; styles 2.5—4 mm long, shortly connate for 0.8—1 mm at base, strongly recurved, papillose adaxially, with dense trichomes abaxially for entire length. Fruits subglobose, weakly trilobate, 12—16 mm long, 13-18 mm diameter, minutely muriculate; seeds 5—9 mm long, 3.5—6 mm wide, 1.5—2 mm thick, smooth, shiny brown. Fig. 3.
Selected additional specimens examined: Queensland. North Kennepy District: Hook Island, Nov 1985, Batianoff 3601A (BRI); Dugong Inlet, Whitsunday Island, 20°15’S, 148°57’E, Sep 1990, Batianoff 900973 & Batianoff (BRD); Walkers Creek, Mt Elliot, S of Townsville, 19°35’S, 146°59"E, Aug 1991, Bean 3617 (BRI); Station Hill, Cape Upstart Headland, 19°46’S, 147°49°E, Sep 1991, Cumming 11376 (BRI); Dryander Creek, left branch, SE slopes of Mt Dryander, 20°16’S, 148°35’E, Jun 1989, Forster 5128 & Tucker (BISH, BRI, MEL, MOQ); Dryander Creek, middle branch, SE base of Mt Dryander, 20°16’S, 148°35’E, Jun 1989, Forster 5175 & Tucker (BISH, BRI, DNA, K, QRS); Emmett Creek, Bowling Green Bay N.P., 19°27’S, 147°03’E, Dec 1993, Forster 14307 (BRI, MEL, QRS); ditto, Feb 1994, Forster 14883 & Bean (A, BRI, L, MEL, QRS); Cape Upstart, 19°42’S, 147°45’E, Oct 1985, Ade/vi/le 592 (BRI); Dryander Creek, c. 2 km N of Gregory & c. 20 km N of Proserpine, 20°24’S, 148°35’E, Nov 1985, Sharpe 4175 (BRI); SE foothills of Mt Dryander, May 1969, Siith 14526 (BRI, CANB).
Distribution and habitat: D. indistincta 1s restricted to the North Kennedy district of Queensland where it occurs in three 1° grid cells between Townsville and Proserpine (Map 2). Plants grow on alluvium and talus along creeks in complex notophyll to mesophyll vineforests over granite substrates.
Notes: D. indistincta may be distinguished from all other species in the genus by the very poorly developed lateral venation in the leaves where only the intramarginal veins are raised on the lower leaf surface. D. indistincta, D. tuckeriand D. laxinervis are closely allied and superficially similar species (Table 1). D. indistincta differs from D. tuckeri in its smaller, deltoid-lanceolate stipules that leave a stipular scar 0.5—0.8 mm wide, greater number of lateral veins in the leaf lamina (8—10) that are also more poorly developed, and shorter styles
19
(2.5—-4 mm long). D. indistincta differs from D. laxinervis in the colour of its new foliage (bright pink-brown), generally shorter petioles, the shorter styles (2.5—4 mm long) that are strongly recurved and shortly connate (for 0.8-l mm), and its male flowers with fewer stamens (16-18) and shorter anthers (0.8—1.3 mm long).
Phenology: Flowers February to June; fruits several months later. Flowering is extremely erratic and appears to be stimulated by heavy rainfall in the summer wet season.
Conservation status: D, indistincta 1s very common where it occurs and is known to be present in National Parks at Mt Elhot and Cape Upstart. No conservation coding is thought necessary.
Etymology: The specific epithet is derived from the Latin indistinctus (indistinct) and alludes to the poor development of lateral veins in the leaf lamina of this species.
4, Dissiliaria tuckeri P.[.Forst., sp. nov. affinis D, laxinervi Airy Shaw a qua frondescentia nova clara rosei-rubra, stipulis magnioribus lanceolati-obovatis (2,.2—-4 mm longis) cicatrice 1.8—2.2 mm lata, et stylis brevioribus (4—4.5 mm longis) valde recurvatis differt. Typus: Queensland. Cook Districr: Gap Creek, c, 22 miles [36.7 km] SE of Cooktown, May 1969, L.S. Smith 14419 (holo: BRI; iso: NSW),
Illustration: Christophel & Hyland (1993: 102, t. 40E); Cooper & Cooper (1994: 97) [as D. laxinervis\.
Tree to 30 m high. Trunk buttressed at base, rarely multistemmed, not suckering; bark rough, flaky in patches, blaze pink. Branchlets + rounded, glabrous or with a few sparse trichomes when young, lenticellate with age. Stipules lanceolate-obovate, 2.2—-4 mm long, 1.8—2.2 mm wide, glabrous. Leaves glabrous, new foliage usually bright red-pink; petioles 4—8 mm long, 1.5—2.5 mm wide, channelled on top, glabrous; lamina elliptic, oblanceolate, 60—170 mm long, 30-75 mm wide, coriaceous, margins entire, venation composed of 8-10
20
lateral veins per side of midrib and weakly developed interlateral reticulate tertiary veins with only the outer veins raised; upper surface dark glossy green, venation weakly developed; lower surface matt light green, venation weakly developed; tip acute, acuminate, obtuse, rounded; base cuneate or rounded. Male inflorescences and flowers not seen. Female inflorescences in distal axils; female flowers | or 2 per axil with peduncles 2—4 mm long; bracts not seen, Female flowers (not seen at anthesis) with pedicels 15-33 mm long, glabrous; styles 44.5 mm long, connate for c. | mm at base, strongly recurved, papillose adaxially top and with sparse trichomes abaxially for entire length. Fruits subglobose, weakly trilobate, 13-15 mm long, c. 18 mm diameter, minutely muriculate; seeds c. 9 mm long, 7 mm wide and 1.5—2 mm thick, smooth, shiny brown. Fig. 4.
Austrobaileya 5(1): 9-27 (1997)
Selected additional specimens examined: Queensland. Cook District: Noah Creek, 3 km upstream from road, 16°08’S, 145°25°E, Aug 1987, Christophel [QRS 084689] (QRS); Gap Creek, Cedar Bay N.P., 15°49’S, 145°19°, Jun 1992, Forster 10729 et al. (BRD); Little Cooper Creek, Datntree Freehold Rainforest, 16°10’S, 145°24’E, Jul 1994, Forster 15558 (BRI, QRS); Cape Tribulation N.P., Noah Creek, 16°08’S, 145°25’E, Sep 1991, Gray 5315 (QRS); 13.2kmS of Rossville on Bloomfield road, lowlands below Mt Finnegan, Dec 1989, Jessup GJD3086 et al. (BRI); Daintree, Nov 1959, Manthey 1777 (BRI); Gap Creek, 15°45’°S, 145°10’E, May 1969, Smith 14372 (BR); Bloomfield, 15°57’S, 145°20’E, May 1969, Volck 4290 (BRI); The Gap, between Rossville & Ayton, Jun 1973, Webb & Tracey 13709 (BRI).
Distribution and habitat: D. tuckeriis known from several disjunct populations in two 1° grid squares in the ‘Wet Tropics’ of Cook district of north Queensland (Map 1). Plants grow in complex mesophyll vineforest on rocky alluvia derived from granite substrates along permanent watercourses.
4 tba t aust Hats 5 t 2 co aay a 1,4 | ¥ ¥ 4 ¥ a x ’ 1 2! F ee # = « 4 a = ag *% a? * | * ws a, *4t vn, : Eo = aes i » t * ahr £, fire r ‘ = a ‘ + Fal utd = fea pe ™ F = +n = é Ace nam, an’ = = Ne or re ba Pa Pa. a,Fu at = - f TER a - ef
Fig. 4. Dissiliaria tuckeri. A. twig x 0.5. B. stipule x 5. C. apical view of fruit x 1.5. D. lateral view of fruit x 1.5. E. lateral view of seed x 2.5, F, ventral view of seed x 2.5. A,B from Smith 14419 (BRI); C-F from Gray 5315 (QRS).
Del. W. Smith.
Forster, Dissiliaria
Notes: D. tuckeri is closely allied to D. laxinervis and the two species may be difficult to distinguish with incomplete herbarium material (see Table 1 for comparison). D. laxinervis has green to greenish pale pink new foliage; 1—1.5 mm long deltoid-lanceolate stipules that leave a scar 0.5—0.8 mm wide, generally shorter leaves (45-130 mm long), and longer styles (c. 6 mm long) that are held erect. D. tuckeri has bright pink-red new foliage; 2.2—4mm long lanceolate-obovate stipules that leave a scar 1.8—2.2 mm wide; generally longer (mainly oblanceolate) leaves (60-170 mm long) and shorter styles (44.5 mm long) that are strongly recurved. There also appear to be some differences in the seedlings of these two species (based on limited data). Seedlings of D. tuckeri have cotyledons that are more or less sessile and 22—-23 mm long, whereas D. laxinervis has cotyledons that are shortly petiolate (c. 2 mm long) and 26-30 mm long.
The populations of D. tuckeri at the type locality of Gap Creek have much larger stipules than those from the Daintree area, but whether this is correlated with other morphological differences remains to be seen. There appear to be some differences in seedlings from the two areas (G. Sankowsky, pers. comm. 1994) but the evidence remains inconclusive at this stage. To determine whether or not D. tuckeri encompasses two taxa requires examination of fertile collections of male and female flowers! These have proved impossible to procure to date.
Conservation status: D. tuckeri is present in Cedar Bay National Park at the foothills of Mt Finnegan (Gap Creek) and in Cape Tribulation National Park at Noah Creek, but is still unreserved at Cooper Creek. The Cooper Creek population 1s under immediate threat due to residential development in one of the most diverse and species-rich areas of lowland rainforest north of the Daintree River. A conservation coding of 2VC as defined by the Queensland Nature Conservation Act 1992 1s recommended.
Etymology: The specific epithet honours Maurie C. Tucker of Booval, Ipswich who has a keen interest in Australian rainforest plants
21
and has explored many patches of rainforest and scrub with the author.
5, Dissiliaria muelleri Baill., Adansonia 7: 359 (1867). Type: Queensland. Port Curtis District: Rockhampton, 7hozet (lecto [here designated]: P. v.v., photo at BRI).
Illustration: Baillon (1867: plate 1).
Tree to 10 m high. Trunk not buttressed, not suckering or only if damaged, but often multistemmed. Bark rough, flaky in patches, light to rich brown; blaze pink. Branchlets -+ rounded, with short sparse trichomes when young, soon glabrescent and lenticellate with age. Stipules lanceolate, 0.8—1.4 mm long, 0.4—0.6 mm wide, glabrous or with scattered trichomes. Leaves glabrous; petioles 1-4 mm long, c. 0.5 mm wide; laminas broadly ovate to rhombic-ovate, 8-95 mm long, 6-65 mm wide, chartaceous, with prominent yellow venation comprising 7 to 9 laterals per side of midrib and weakly developed interlateral reticulate veins; upper surface glabrous, glossy green, venation + obscure; lower surface glabrous, pale green, venation prominent; margins shallowly crenulate to subentire, with 14-16 teeth per side; tip acute, obtuse or retuse; base cuneate, obtuse, truncate. Male inflorescences 1—few-flowered in distal axils; peduncles !~3 mm long, glabrous; bracts lanceolate to ovate, 1.4—2 mm long, 0.5—-1.8 mm wide, olabrous. Male flowers with pedicels 2.5—7 mm long, c. 0.4 mm diameter, glabrous; sepals broadly elliptic, 1.5-2 mm long, 1.4—-1.5 mm wide, glabrous or ciliate; receptacle c. 2 mm diameter, with sparse to dense trichomes and numerous small glands; stamens 8-16; filaments filiform, 0.9—1.8 mm long, c. 0.1 mm diameter; anthers 0.5—-1 mm long, 0.3—-0.8 mm wide. Female inflorescences (1-)—3-flowered, up to 15 mm long; peduncles 4-10 mm long, 0.7-0.8 mm diameter, glabrous; bracts lanceolate to oblanceolate, 0.8—1.5 mm long, 0.3-0.8 mm wide, glabrous. Female flowers with pedicels 1.5-10 mm long, 0.7—1.2 mm diameter, glabrous; sepals broadly elliptic to ovate, 1.8-7.5 mm long, 1.2—-3 mm wide,
glabrous; disk annular to cupular, surrounding
the base of the ovary; ovary subglobose, 1.3-4 mm diameter, with sparse trichomes; styles very shortly connate, subulate, erect to
22
slightly recurved, 2—3.2 mm long, with sparse trichomes at base. Fruits subglobose, strongly trilobate, 8-9 mm long, 10-15 mm diameter, with scattered to sparse trichomes, yellow-green. Seeds 4.8—5.5 mm long, 3.7-4.5 mm wide, 22.2 mm thick, smooth, shiny red-brown. Fig. 5.
Selected additional specimens examined: Queensland. Port Curtis District: Rockhampton, Dallachy [MEL 532391] (MEL); Mt Larcom, 23°48’S, 151°04’E, Jan 1988, Forster 3372 & Gibson (BRI, QRS); ditto, Jan 1994, Forster 14640 (BRI, MEL, QRS); Moores Creek E.P., Beserker Range, 23°19’S, 150°33’E, Jan 1993, Forster 12713 (BRI, L, MEL, QRS); Moores Creek, 23°20’S, 150°3S5’E, Oct 1976, Hyland 9084 (BRI, QRS); Archer Creek near Rockhampton, Jul 1935, White 12142 (BRI). Wipe Bay Districr: Graphite Mine road, Guyra Range, Mt Bauple area, 25°47°S, 152°34’E, Dec 1992, Forster 12519A & Smyre/l (BRL K, L, MEL, QRS), 12519B (BRI, K, L, MEL, QRS); ditto, Jan 1994, Forster 14629 (BRD; c. 1 km N of Mt Bauple, 25°49’°S, 152°35’E, Dec 1982, Guymer 1834 & Dillewaard (BRI); Mt Bauple, 27 miles [45 km] N from Gympie, Dec 1923, Kajewski[AQ20228 1] (BRD); Gundiah, Dec 1923, Kajewski [AQ202279] (BRI).
Distribution and habitat: D. muelleriis known from three localities, Mt Archer near Rockhampton, Mt Larcom and Mt Bauple (Map 2), all in central and south-east Queensland (Forster et al. 1991), Plants grow in semi-evergreen microphyll vinethickets and araucarian microphyll vineforests on volcanic substrates of volcanic orgin.
Notes: Baillon cited two syntypes for D. muelleri, Thozet391 and Dallachy 429, both collected from the Rockhampton area. I have been able to locate unnumbered collections from the Rockhampton area of both these collectors in MEL and P. The Thozet collection at P is the better collection and hence is selected as lectotype of the name.
Sterile collections from the Mcllwraith Range area in Cook district have been referred to this species in the past (e.g. Airy Shaw 1981); however, these belong to a distinct taxon named D. surculosa in this paper.
Phenology: Flowers from December to January; fruits January to February.
Conservation status: D. muelleriis a restricted endemic that is locally abundant in the three general localities where it occurs. Parts of two
Austrobaileya 5(1); 9-27 (1997)
of these populations are in conservation reserves (Mt Bauple National Park & Mt Archer Conservation Park), but the Mt Larcom population is unreserved. No conservation coding 1s considered necessary at this stage.
6. Dissiliaria surculosa P.I.Forst., sp. nov. affinis D. muelleri Baill. a qua caulibus juvenibus dense lenticellatis, foliis anguste ellipticis et dentibus utroque latere 17—23 et apice acuminato, et sepalis florum marium obovatis et 0.8—1.2 mm latis differt. Typus: Queensland. Coox District: Timber Reserve [4 Massy, 13°52’S, 143°23’E, 8 Nov 1980, B. Hyland 10882 (holo: QRS;
iso: BRI).
Dissiliaria sp. RFK#2554 (Hyland & Whiffin 1993, 2: 132).
Dissiliaria sp. (Rocky River B.Hyland 10882) (Forster 1994).
Illustration: Christophel & Hyland (1993: 102F).
Tree up to 13 m high. Trunk not buttressed, often suckering strongly from base. Bark brownish, with numerous dark brown pustulate lenticels; blaze pink. Branchlets + rounded, with sparse trichomes when young, glabrescent, with dense lenticels even when young. Stipules lanceolate, 2~—2.5 mm long, c. 0.7 mm wide, with sparse trichomes. Leaves with sparse trichomes when young, glabrescent, pale green; petioles 1.5-3 mm long, 0.5—0.8 mm diameter; laminas narrowly elliptic, 22~65 mm long, 7—27 mm wide, chartaceous, venation composed of 7 or 8 prominent lateral veins per side of midrib and interlateral reticulate tertiary veins; margins crenate with 17 to 23 teeth per side of midrib; upper surface glossy green, venation + obscure; lower surface pale green, venation prominent; tip short to long acuminate; base cuneate. Male inflorescences 1~—3 in distal axils with peduncles up to 2 mm long, with sparse trichomes; bracts lanceolate, 1—1.8 mm long, 0.30.4 mm wide, glabrous. Male flowers with pedicels 5-10 mm long, c. 0.3 mm diameter, with scattered trichomes at base; sepals obovate, 1.8—2.3 mm long, 0.8—1.2 mm wide, glabrous or with scattered trichomes and
Ma eee
Petree eet ee tert!
Forster, Dissiliaria 23
a? x
A?
om =F =4
Fig. 5. Dissiliaria muelleri. A. twig x 1, B. detail of adaxial leaf surface x 2. C. female inflorescence < 8. D. male inflorescence = 8. E, male flower x 16. F. stamen x 32. G. apical view of fruit x 4. H. lateral view of fruit x 4, I. lateral view of seed x 8. J. ventral view of seed x 8. A,B from Forster 12519A & Smyrell (BRI); C from Forster 12713 (BRI); D-F from Forster 12519B & Smyrell (BRD); G,H from Forster 3372 & Gibson (BRI); I,J from Forster 14629 (BRI). Del. W. Smith.
24
marginal cilia; disk c. 1.6 mm diameter, with dense trichomes; stamens 16; filaments filiform, 1—-1.4 mm long; anthers 0.8—1 mm long, 0.3—-0.5 mm wide. Female inflorescences and flowers not seen. Fruit subglobose, strongly trilobate, 9-10 mm long, 11-12 mm diameter, + glabrous. Seed not seen. Fig. 6.
Selected additional specimens examined, Queensland. Cook District: Temple Creek Catchment, Cape Melville N.P., 15 km NW of Barrow Point, 14°17°S, 144°31’E, Sul 1993, Fell 3331 et al (BRI, CANB, DNA, MBA, MEL, QRS); Rocky River Scrub, Silver Plains Station, 13°48’S, 143°28’E, Jun 1992, Forster 10606 et al (BRI, QRS); ditto, Jul 1993, Forster 13620 et al. (BRI, QRS); Upper Massey Creek, c. 15 miles [25 km] ENE of Coen, Oct 1962, Smith 11730 (BRI); MclIlwraith Range, c. 11 miles [18.3 km] ENE of Coen, Oct 1962, Smith 11792 (BRD; Rocky River, eastern foothills of McIlwraith Range, 13°47°S, 143°25’E, Oct 1969, Webb & Tracey 9435 (BRI).
E
as a te
5
CAPRIS | Seer anitee pat SE EEE HEP? H HY
Austrobaileya 5(1): 9-27 (1997)
Distribution and habitat. D. surculosa 1s endemic in Cape York Peninsula where it is known from the eastern fall of the MclIlwraith Range and Cape Melville (Map 2). At the Mcliwraith Range, plants grow near watercourses in semideciduous notophyll to mesophyll vineforest on deep alluvia derived from metamorphic and granite substrates. This gallery-forest is usually flooded during the wet season and in July 1993, there was evidence of flood debris some 12 m above the ground in the crowns of the trees. Associated plants include Castanospermum australe A.Cunn. & Fraser ex Hook., Ficus albipila (Miq.) King, Haplostichanthus sp. (Rocky River Scrub P.].Forster PIF10617), Rhodamnia australis A.J. Scott, Pilidiostigma recurvum (C.T. White) A.J.Scott, and Codiaeum variegatum var. moluccanum (Decne.) Muell.Arg. At Cape Melville, the plants grow in semi-deciduous
anvae
Map 2. Distribution of @ Dissiliaria indistincta, A D. surcilosa and * D. muelleri
Forster, Dissiliaria 25
Pa ¢
7 Seer
f at / tts
he af. + al ai ‘oat eaiteri, fay
Mess Cad
bh ‘ane!
“ hI hd gt aft hat et Fatal
at +t
gat
La
q me Sth
ot aeetis ote
; ¢P athe
Fig. 6. Dissiliaria surculosa. A. twig. x 1. B. node with male inflorescence x 2.5. C. male flower x 5, D. stamen x 10. E. apical view of fruit x 2, F, lateral view of fruit x 2. G. ventral view of seed x 5. H. lateral view of seed x 5. I. seedling x 1, A-D from Hyland 10882 (BRI); E-I from Forster 13620 (BRI). Del. W. Smith.
SRE nye inal aug eo Sah oh lagi Bones one sae
26
notophyll to microphyll vinethicket on hillslopes on colluvium derived from granite. Associated canopy species at this locality include Pongamia pinnata (L.) Pierre and Wodyetia bifurcata Irvine.
Notes: D. surculosa appears to have been first collected by L.S.Smith in 1962. Sterile material of this plant was referred with doubt to D. muelleri by Airy Shaw (1981).D. surculosa strongly resembles D. muelleri, but differs in its narrowly elliptic leaf laminas with a short to long acuminate tip, the 17—23 marginal teeth per side of the midrib, the dense lenticels on young stems and the sepals of the male flowers being obovate and 0.8—1.2 mm wide.
Conservation status: D. surculosa is apparently restricted in its distribution. At present there appear to be no obvious threats to this plant and it is present in Cape Melville National Park. No conservation coding is thought necessary.
Etymology: The specific epithet is derived from the Latin surculus (shoot, sucker) and-osa (abundant) which alludes to the tendency of this plant to profusely sucker from the stems. This suckering could be due to damage from irregular flooding in the habitat.
Excluded species
l. Dissiliaria tricornis Benth., Fl. Austral. 6: 91 (1873).
This is Choriceras tricorne (Benth.) Airy Shaw.
2. Dissiliaria sp. RFK#25730 (Christophel &.
Hyland 1993: 102g; Hyland & Whiffin 1993, 2: 132).
Dissiliaria sp. (Rex Range, G.Sankowsky 1075) (Forster 1994).
This was described as Sankowskya stipularis P.1.Forst. (Forster 1995).
Acknowledgements
The illustrations were drawn by W.Smith (BRI) with partial funding from the Australian Biological Resources Study (ABRS). Fieldwork and/or special collections were made with the
Austrobaileya 5(1): 9-27 (1997)
assistance of A.R.Bean, D.G.Fell, D. & [.Liddle, P.Machin, G.Smyrell, G. & N.Sankowsky and M.C.Tucker. Photographs of material at BM and K were organised by P.S.Short (MEL) while Australian Botanical Liaison Officer (ABLO) at Kew (U.K.). Photographs of material at P were taken by G.P.Guymer (BRI) while ABLO at Kew (U.K.). The diagnoses were translated into Latin by L.A.Craven (CANB). Comments on a draft of the manuscript were provided by A.R.Bean (BRI). The Directors/Curators of the cited herbaria allowed access to collections either on loan or in sifu. B. Hyland and staff at QRS provided support and access to facilities on repeated visits to north Queensland. Access to populations of Dissiliaria species in State Forests and Timber Reserves was facilitated by permits from the Queensland Forest Service, Department of Primary Industries. The work was funded as a preferred objective by ABRS during 1992-1994. Additional fieldwork in the ‘Wet Tropics’ region of north-east Queensland was possible due to a travel grant from the Wet Tropics Management Authority in 1993-1994 for work on Endangered Euphorbiaceae in that region.
References
Airy SHAW, H.K. (1980), New or noteworthy Australian Euphorbiaceae - I. Adwelleria 4: 207-241.
———- (1981). A partial synopsis of the Euphorbiaceae - Platylobeae of Australia (excluding Phyllanthus, Euphorbia and Calycopeplus). Kew Bulletin 35: 577-700,
BAILLoN, H.E. (1867). Species Euphorbiacearum. Recherches complementaires sur les Euphorbiacées Australiennes, Adansonia 7: 352-360,
Bartow, B.A. & HyLanp, B.P.M. (1988). The origins of the flora of Australia’s wet tropics. Proceedings of the Ecological Society of Australia 15: 1-17.
BEnTHAM, G. (1873). Euphorbiaceae, In Flora Australiensis 6: 41-153. London: L. Reeve & Co.
CuapMAN, A.D. (1991), Australian Plant Name Index. Canberra: Australian Government Publishing Service,
CHRISTOPHEL, D.C. & Hy.anp, B.P.M. (1993). Leaf Atlas of Australian Tropical Rain Forest Trees. Melbourne: CSIRO Publications.
Cooper, W. & Cooper, W.T. (1994). Fruits. of the Rainforest. Chatswood (Sydney): RD Press.
PUSS ESN Sen Sg ect ocionicennxsneeinideddieho
“ISTE
Forster, Dissiliaria
Forstser, P.I. (1994). Euphorbiaceae (in part). In R.J.F.Henderson (ed.), Oueensland Vascular Plants: Names and Distribution, Brisbane: Queensland Department of Environment & Heritage.
— (1995). Sankowskya, a new genus of Euphorbiaceae (Dissiliariinae) from the Australian Wet Tropics. Austrobaileya 4; 329-335.
Forster, P.l., Bostock, P.D., Biro L.H. & BEAN, A.R. (1991). Vineforest Plant Atlas for South-east Queensland, Brisbane: Queensland Herbarium.
Francis, W.D. (1981). Australian Rain-forest Trees, 4th Ed. Canberra: Australian Government Publishing Service.
Hauser, J. (1992). Fragments of Green. Bardon (Brisbane): Rainforest Conservation Society Inc.
Haypen, W.J. (1994). Systematic anatomy of Euphorbiaceae subfamily Oldfieldioideae. I. Overview. Annals of the Missouri Botanical Garden 81: 180-202.
Hewson, H. (1988), Plant Indumentum. A Handbook of Terminology. Austraitan Flora and Fauna Series No. 9. Canberra: Australian Government Publishing Service.
HYLAND, B.P.M. & WuirFin, T. (1993). Australian Tropical Rain Forest Trees: An Interactive Identification System. Melbourne: CSIRO Publications.
Levin, G.A, & Simpson, M.G. (1994), Phylogenetic implications of pollen ultrastructure in the Oldfieldioideae (Euphorbiaceae). Annals of the Missouri Botanical Garden 81: 203-238.
Rao, P.N. & Ragu, V.S. (1985). Foliar trichomes in the family Euphorbiaceae. In C.M. Govil & V. Kumar (eds.), Trends in Plant Research, pp. 128-136. Dehra Dun: Bishen Singh & Mahendra Pal Singh (n.v., cited by Hayden 1994),
Wess, L.J. (1978). A general classification of Australian rainforests. Australian Plants 9: 349-363.
Wess, L.J. & Tracey, J.G. (1981). Australian rainforests: pattern and change. In A. Keast (ed.), Ecological Biogeography of Australia. pp. 605-694, The Hague: W. Junk.
Wester, G.L. (1994). Synopsis of the genera and suprageneric taxa of Euphorbiaceae. Annals of the Missouri Botanical Garden 81: 33-144.
27
aT CT ae eT ea nN eee nen Re ML DOR SL ce DLC ace enc Cir COC rare Gtr us rear eee a eas re ee
EIS Tn ea Se A De SI
Three new species of Fontainea Heckel (Euphorbiaceae) from Australia and Papua New Guinea
Paul I. Forster
Summary
Forster, Paul I. (1997). Three new species of Fontainea Heckel (Euphorbiaceae) from Australia and Papua New Guinea. Austrobaileya 5(1): 29-37, Fontainea fugax P.I.Forst. from Queensland, Australia, and F’. borealis P.I.Forst. and F. subpapuana P.1.Forst. from Papua New Guinea are named as new species. Information is provided on their distribution, habitat and affinities. Separate identification keys are provided for Fontainea in New Guinea and Australia.
Keywords: Euphorbiaceae, Fontainea, Fontainea borealis, Fontainea fugax, Fontainea subpapuana,
Papua New Guinea, Queensland.
Paul I. Forster, Queensland Herbarium, Meiers Road, Indooroopilly, Queensland 4068, Australia
Introduction
The genus Fontainea Heckel is classified in Euphorbiaceae, subfamily Crotonoideae, tribe Codiaeae (Webster 1994) and comprises shrubs or small trees that grow in rainforest communities. Jessup & Guymer (1985) revised Fontainea and recognised six species for Australia and New Caledonia, as well as mentioning additional unnamed taxa from Papua New Guinea and New Caledonia.
Airy Shaw (1974, 1980) referred the collections of Fontainea from Papua New Guinea to F. pancheri (Baillon) Heckel based on their similarity to the type of this species from New Caledonia. Jessup & Guymer (1985) thought this identification unlikely, but did not resolve the identity of the species in Papua New Guinea due to the incomplete nature of the available specimens.
In preparing accounts of Fontainea, both for ‘Flora of Australia’ and ‘Flora Malesiana’, it is now necessary to determine the status of the Fontainea collections from Papua New Guinea. Although no further collections have been made in the decade since Jessup & Guymer’s account, the available collections do enable diagnostic characters to be determined, as well as preparation of descriptions and discussion of affinities. There are two species
Accepted for publication 25 July 1997
of Fontainea present in Papua New Guinea, one from the north-east and one from the south- east parts of the country. These are named as F, borealis and F. subpapuana_ respectively. An additional species of Fontainea from south- east Queensland, Australia has also been discovered tn the last decade, and is named FF’ fugax.
Taxonomy
1. Fontainea borealis P.I.Forst., sp. nov. affinis F’, picrospermae C,T.White a qua lamina folu prope basem eglandulosa, calyce 4—lobato lobatis ovatis, petalis obovatis usque lanceolatiovatis, floribus maribus disco c. 0.6mm alto lobato irregulariter, et staminibus connatis 1—1.5 mm super discum differt.Typus: Papua New Guinea. EASTERN HIGHLANDS PROVINCE: Aiyura, Oct 1944, LS. Smith NGF1030 (holo: K; iso: BRI; L n.v.).
Shrub or small tree to 12 m high; stem exudate colour unknown. New shoots with dense, antrorse golden trichomes. Stipules absent. Leaves petiolate, discolorous; petioles 10-26 mm long, 1—1.4 mm diameter, swollen at base and apex, narrowly channelled above, glabrous; lamina elliptic to oblanceolate, 56-240 mm long, 22-70 mm wide, not decurrent, chartaceous; upper surface dark green; lower surface pale green; venation comprising 8—14 lateral veins per side of midrib and reticulate interlateral veins; tip acute to
30
acuminate; base cuneate; glands absent from base of lamina. Male inflorescences terminal or axillary, with sparse trichomes. Male flowers 5—7 mm long, 7-8 mm diameter; pedicels 2.5—-5 mm long, c. | mm wide, with scattered trichomes; calyx 4-lobed, 2.8-3.5 mm long,
lobes ovate; petals 5, lanceolate-ovate to obovate, 5—6 mm long, 2~3.5 mm wide, weakly
recurved, externally and internally with dense velutinous trichomes; disk undulate with srooves, c. 0.6 mm long and 2.5 mm diameter, glabrous; stamens 24-28, connate 1—1.5 mm above top of disk, filaments free for 2—2.5 mm,
Austrobaileya 5(1): 29-37 (1997)
glabrous towards top, densely hairy at base; anthers c. 0.8 mm long and 0.5 mm wide. Female flowers not seen. Fruit not seen. Fig. 1.
Additional specimens examined: Papua New Guinea. EASTERN HIGHLANDS PROVINCE: Numura, an Strasse Kainantu - Goroka, 0.5 mile nach Abzweigung nach Okapa, 6 mile von Kainantu, Jan 1964, Stauffer 5608 & Sayers (K; L 2.V.).
Distribution and habitat: Fontainea borealis is known only from the Eastern Highlands Province of Papua New Guinea. Plants have been recorded from rainforest at altitudes of 1800—2000 m.
Notes: The available herbarium material of F’. borealis 1s incomplete thus preventing a detailed description. F. borealis appears to be allied to the Australian F’. picrosperma C.T.White and a comparison of these two taxa is made in Table 1.
Table 1. Morphological comparison of Fontainea picrosperma and F. borealis.
Character FP’, picrosperma F. borealis
glands in leaf lamina present absent
disk undulate to lobed iregularly lobed 0.7—1 mm high c. 0.6 mm high
male calyx 2—3-lobed 4-lobed
male calyx lobes triangular-ovate to ovate
male petals
stamens connate c. 0.5 mm
oblong-ovate
broadly ovate
obovate to lanceolate-ovate
1—-1.5 mm
Forster, Three new species of Fontainea Heckel
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Conservation status: Unknown.
Etymology: The specific epithet is derived from Latin borealis (northern) and refers to the northerly distribution of this species within the genus.
2. Fontamea subpapuana P.I.Forst., sp. nov. affinis F. picrospermae C.T.White a qua petiolo tumido ad apicem tantum, lamina folii venis primariis utroque costate 13-15, calyce florum marium 4—5—lobato, stylis longioribus (33.5 mm longis), et endocarpo paginis intersuturalibus latioribus (12-13 mm latis) differt. Typus: Papua New Guinea. CENTRAL PROVINCE: Kuriva Forestry area, near Weimauri River, 9°05’S, 147°05’E, 6 May 1971, H.Streimann & A.Kairo LAES51548 (holo: K; iso: BRI, K; Lv.v.).
Tree to 7 m high; stems with red exudate. New shoots with dense, antrorse yellow trichomes. Stipules absent. Leaves petiolate, discolorous; petioles 12—20 mm long, 1-1.4 mm diameter, swollen at top and narrowly channelled above, with scattered trichomes; lamina elliptic to oblanceolate, rarely obovate, 60-170 mm long, 30-60 mm wide, not decurrent, chartaceous; upper surface dark green; lower surface pale green; venation comprising 13—15 lateral veins and reticulate
Austrobaileya 5(1): 29-37 (1997)
interlateral veins; tip acuminate; base cuneate to weakly attenuate; glands elliptic, sessile, I—2 mm long,+ marginal, 0.5—lmm from base oflamina. Male inflorescences not seen. Male flowers not seen. Female inflorescence terminal, comprising 1—3 flowers. Female flowers 5—8 mm long, 10-13 mm diameter; pedicels 7-15 mm long, I~1.2 mm diameter, with sparse trichomes; calyx 4—5-lobed, 2—2.5 mm long, lobes rounded-ovate, with sparse trichomes; petals 5, lanceolate-ovate, 6-8 mm long, 2.5—3 mm wide, + recurved, with dense velutinous trichomes externally and internally; disk not dissected; ovary 3—4-locular, ovoid, with dense antrorse trichomes; styles 3—3.5 mm long. Intact fruit not seen. Sarcocarp red (n.v.). Endocarp very shortly beaked, 3-ridged at sutures; mintersutural faces smooth, convex, 25-26 mm long, 12-13 mm wide. Fig. 2.
Distribution and habitat: Fontainea subpapuana 1s known only from Central Province in south-east Papua New Guinea. Plants were recorded as growing in lowland rainforest on river flats.
Notes: Fontainea subpapuana is very poorly known and the above description lacks significant details with regard to the male flowers and mature fruit. This species is closely allied to the Australian F’. picrosperma. Fontainea subpapuana and F. picrosperma are compared in Table 2. Fontainea subpapuana and F. borealis may be distinguished with the following key.
1. Leaf petioles swollen at base and apex; glands absent from
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34
Austrobaileya 5(1): 29-37 (1997)
Table 2. Morphological comparison of Fontainea picrosperma and F. subpapuana.
Character
No. lateral veins per 8—12
side of leaf lamina midrib
Petiole base swollen Petiole apex swollen male flower calyx 2—3-lobed style length (mm) 0.8—2 endocarp intersutural face
width (mm) 7-10
Conservation status: Unknown.
Etymology: The specific epithet is derived from Latin sub- (below) and papuana (of Papua), and refers to the distribution of this species on the south coast of Papua New Guinea.
3. Fontainea fugax P.I.Forst., sp. nov. affinis F. rostratae Jessup & Guymer a qua petiolo ad basem etumido, floribus maribus minoribus (4-5 x 6—8 mm) et staminibus paucioribus (24), endocarpo rostro breviore (1—1.7 mm longo) et paginis intersuturalibus leniter rugosis plus minusve planis usque leniter convexis et perminore (15-17 x 89.5 mm) differt. Typus: Queensland. Burnett District: Fontainea Scrub, State Forest 172, Gurgeena Plateau, 9 Feb 1994, PI. Forster 14802 (holo: BRI {1 sheet + spirit, male plant])
Shrub to 4 m high; stems with clear exudate. New shoots with sparse, antrorse uncoloured trichomes. Stipules absent. Leaves petiolate, discolorous; petioles 2—4 mm long, c. 1 mm
fF. picrosperma
F’, subpapuana
13-15
not swollen
swollen
4—5-lobed
3—3.5
12-13
diameter, of uniform thickness and narrowly channelled above, glabrous; lamina elliptic, obovate or oblanceolate, 15—75 mm long, 6—42 mm wide,+ decurrent, chartaceous; upper surface dark green; lower surface pale green; venation comprising 6—10 lateral veins and reticulate interlateral veins; tip acute to obtuse; base cuneate; glands elliptic, sessile, 1—2 mm long, markedly intramarginal, 2-8 mm from base of lamina. Male inflorescence terminal or axillary, with sparse trichomes. Male flowers 4—5 mm long, 6-8 mm diameter; pedicels 2.5—5 mm long, 0.5—1 mm wide, with sparse trichomes; calyx 4-lobed, 2—2.5 mm long, lobes ovate; petals 4, lanceolate-ovate, 4.5—5 mm long, 2—2.5 mm wide, strongly recurved, externally with sparse trichomes, internally with sparse trichomes apart from near base where they are dense and velutinous; disk irregularly crenate, 0.4—0.8 mm long, 2-3 mm diameter, glabrous; stamens 24, connate up to 1.5 mm above top of disk, filaments free for 1-3 mm, glabrous towards top, densely hairy at base; anthers 0.6—0.8 mm long, 0.4—0.5 mm wide. Female flowers not seen. Intact fruit not seen. Sarcocarp with sparse trichomes. Endocarp
This content downloaded from 136.154.23.253 on Tue, 18 Jul 2023 23:49:57 +00:00 All use subject to https://about.jstor.org/terms
Forster, Three new species of Fontainea Heckel
beaked for 1—1.7 mm at tip, 3—4-ridged at sutures; intersutural faces weakly rugose,+ flat to slightly convex, 15—17 mm long, 8—9.5 mm wide. Fig. 3.
Additional specimens examined: Queensland. BURNETT District: Portion 90, S.F. 172 [now a Conservation Park}, Aug 1988, Forster 4687 (BRI); Fontainea Scrub, S.F. 172, Oct 1993, Forster 14130 (BRI); ditto, May 1996, Forster 19158 (BRI).
Distribution and habitat: Fontainea fugax 1s known only from the Binjour—Gurgeena Plateau. area between Gayndah and Mundubbera in the Burnett district of south- east Queensland. Plants grow as understorey shrubs in semi-evergreen vine thicket with a canopy dominated by Backhousia kingii
35
Guymer on red euchrozem soils at altitudes between 350 and 400 m.
Notes: Fontainea fugax is closely allied to F’. rostrata Jessup & (Guymer and the initial sterile collection of the plant was referred to as this species (Forster et al. 1991). A comparison of important diagnostic differences between F. fugax and F. rostrata is provided in Table 3. This comparison 1s based on limited material of male flowers and endocarps, yet provides clear morphological discontinuities between the two entities. Unfortunately it has not been possible to obtain herbarium material of female flowers and mature fruits, this is despite repeated visits (c. 10) between 1994 and mid 1996 after rainfall events when flowering could be expected. This situation could persist indefinitely and due to the rarity of this plant it is considered more beneficial for its conservation if the species is formally named.
Table 3. Morphological comparison of Fontainea fugax and F. rostrata.
Character FP’. fugax F’. rostrata
petiole base not swollen base slightly swollen male flowers 4—5 x 6-8 mm 4-6 x 11-13 mm stamen no. 24 28—40
endocarp beak
intersutural faces
Conservation status: Fontainea fugax 1s known from only two populations in close proximity to one another. One population (apparently of one plant) is present in a Conservation Park (site 50 of Forster et al 1991). The other population (with less than 50 plants) is present in State Forest within a very small stand of vine thicket that is experiencing repeated fire incursions on its western boundary and weed infestations of
1—1.7 mm long
weakly rugose + flat to concave 15—17 x 8-9.5 mm
2—3 mm long
strongly rugose + flat to weakly convex 20-23 x 11-13 mm
Mexican daisy — Verbesina encelioides (Cav.) Benth. & Hook.f. ex A.Gray and Dutchmans Pipe—Aristolochia elegans Mast., on the eastern boundary associated with fence construction by the leaseholder. Virtually all semi-evergreen vine thicket on the Binjour—Gurgeena Plateau that is not in conservation Park or State Forest, has now been cleared for agriculture and grazing. This area is notable for other endangered or geographically significant plant
This content downloaded from 136.154.23.253 on Tue, 18 Jul 2023 23:49:57 +00:00 All use subject to https://about.jstor.org/terms
36 Austrobaileya 5(1); 29-37 (1997)
species that occur in the semi-evergreen vine 14134). Both F. fugax and the community in thicket communities, especially Pomaderris | which it occurs, are endangered in the central clivicola E.M.Ross, Bertya opponens(F.Muell. Burnett district. A conservation coding of E ex Benth.) Guymer, B. pedicellata F.Muell. (endangered) is recommended for listing under and Zieria sp. (Binjour Plateau P.1.Forster the Queensland Nature Conservation Act 1992.
Etymology: The specific epithet is derived from Latin fugax (fleeting, transitory) and alludes to the difficulty of obtaining material of the flowers of this species,
= a = = A. 72 eee 77% eA ee fe, Seite Soy tit Laks 4 Veg! iF) tee ie i
Fig. 3. Fontainea fugax P.1.¥orst. A. twig x 0.8. B. undersurface of leaf showing venation < 1.5. C. male flower x 6, D-—F. views of endocarps x 2. A-C Forster 14802 (BRI), D-F Forster 14130 (BRI). Del. W. Smith.
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Forster, Three new species of Fontainea Heckel
37
Key to the Australian taxa of Fontainea
1. Leafpetioles slightly swollen at base and apex; male flower disk with dense
CRIGHONIES — 5 crcte tony Wk lene rE a hs
Leaf petioles of uniform thickness or slightly swollen only at base; male
flower disk glabrous ...........0.005-
2. Leaf petioles not geniculate; basilaminar glands marginal; fruit endocarp
FORIRG GT oF Ue Sink ubnd o dint endvasssivany cee Sudan
Leafpetioles+ geniculate; basilaminar glands intramarginal to submarginal;
fruit endocarp not beaked .............
3, Fruit broadly ovoid, 27—27 mm long; endocarp intersutural faces smooth
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Fruit globose to depressed-globose, 18—22 mm long; endocarp intersutural
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© nan hates, aha ioe “dann cd. bua & ELC oa ea GB eile F. oraria
4, Leaf basilaminar glands marginal, at junction of petiole and lamina; endocarp not beaked, intersutural faces smooth with scattered vascular
FOU AIIIMAG rac sca & pia shock fepephasbthdos gla pd wee hheg? a Stciie Belen ta eed ee eater RON eo leg F. venosa Leaf basilaminar glands intramarginal to submarginal, 2—22 mm from lamina base; endocarp beaked for 1-3 mm, intersutural faces rugose .............. 5 5. Leaf petioles of uniform thickness; fruit endocarp beaked for 1—-1.7 mm; endocarp intersutural faces weakly rugose, 15-17 mm long, 8—9.5 mm VTS) org cg Sos Rod CSUR Toe Rh at WHR. HibeNP AR dans i deel alu hace MRA A Ra oeeaien @ & Me F. fugax
Leaf petioles slightly swollen at base; fruit endocarp beaked for 2—3 mm; endocarp intersutural faces strongly rugose, 20-23 mm long, 11-13 mm
wid Dee is! op ie a wh eed Sang WE weld RGR eh fe Sse tee oo oe Set wen: of wets
Acknowledgements
Thanks to W. Smith (BRD) for the illustrations of F’ fugax, P.Robins (BRI) for the photographs of specimens, and L.A.Craven (CANB) for translation of the diagnoses into Latin and the Directors/Curators of K and L for loan/ information of/on specimens. The Queensland Forest Service of the Queensland Department of Primary Industries is acknowledged for permits to collect and traverse in State Forests and Timber Reserves. This work was funded by the Australian Biological Resources Study during 1992-1994,
References
Airy Suaw, H.K. (1974). The genus Fortainea in New Guinea. Kew Bulletin 29: 326-328.
Airy SHAW, H.K. (1980). The Euphorbiaceae of New Guinea. Kew Bulletin Additional Series VIII. London: Royal Botanic Gardens, Kew.
Forster, P.I., Bostock, P.D., Biro, L.H, & BEAN, A.R. (1991). Vineforest Plant Atlas for South-east Oueensland, Brisbane: Queensland Herbarium.
Jessup, L.W. & Guymer, G.P. (1985). A revision of Fontainea Heckel (Euphorbiaceae: Cluytieae). Austrobaileya 2: 112-125.
Wesster, G.L. (1994). Synopsis of the genera and suprageneric taxa of Euphorbiaceae. Annals of the Missouri Botanical Garden 81: 33-144,
A revision of Rubus subg. Malachobatus (Focke) Focke and Rubus subg. Diemenicus A.R.Bean (Rosaceae) in Australia
A.R. Bean
Summary
Bean, A.R. (1997). A revision of Rubus subg. Malachobatus (Focke) Focke and Rubus subg. Diemenicus A.R.Bean (Rosaceae) in Australia. Austrobaileya 5(1): 39-51. The five indigenous and naturalised Australian taxa belonging to Rubus subg. Malachobatus (Focke) Focke and Rubus subg. Diemenicus subg. nov. are treated. A new variety, R. moluccanus var. trilobus A.R.Bean 1s described. A key to the native and naturalised Australian taxa of Rubus is presented,
Keywords: taxonomy, Rosaceae, Rubus, Australian flora, key, Rubus subg. Malachobatus, Rubus subg. Dalibarda, Rubussube. Diemenicus, Rubus alceifolius, Rubus moluccants var. moluccanus, Rubus moluccanus var. trilobus, Rubus hillii, Rubus x novus, Rubus gunnianus,
A.R.Bean, Queensland Herbarium, Meiers Road, Indooroopilly, Queensland, 4068, Australia
Introduction
Rubus L. is a large genus with many representatives in the northern hemisphere, but it is poorly represented in the southern hemisphere. For example, five indigenous species are recorded for New Zealand (Webb et al. 1980). Australia is similarly poorly endowed, with a total of eight indigenous species, one hybrid and some infraspecific taxa (Bean 1995, 1997).
Two subgenera are treated in this paper:
Rubus subg. Malachobatus (Focke) Focke is a large subgenus with an estimated 127 species (Thompson 1997), and with its centre of diversity in eastern Asia. Twenty species have been recorded for Malesia (Kalkman 1984), New Guinea has three indigenous species, and Australia has just one (R. moluccanus L.). A second species, R. alceifolius Poir. is naturalised in north Queensland. Rubus rugosus Sm., while not known to be naturalised mm Australia, is occasionally cultivated, and has been misidentified by Australian botanists as either R. moluccanus (Symon 1986) or R. Aillti (Bennett 1987).
Accepted for publication 25 July 1997
Rubus gunnianus was formerly included by Focke (1910) with four other species (R. lasiococcus Gray, RK. pedatus Sm., R. fockeanus Kurz and R. dalibarda L.) in Rubus subg. Dalibarda (L.) Focke. The first three of these species were transferred to R. subg. Cylactis (Focke) Focke by Bailey (1941), and all botanists since Focke have accorded R. dalibarda generic status, as Dalibarda repens L. (e.g. Batley 1941, Gleason 1952, Scoggan 1978). R. gunnianus has a number of unique characteristics (see discussion later) and a new subgenus, Rubus subg. Diemenicus A.R.Bean 1s erected here for it.
This is the third and final paper revising the Australian species of Rubus. It is not intended to deal with the taxonomy of naturalised taxa of Rubus L. subg. Rubus, and in the following key, the European Blackberry complex (A. fruticosus agg.) 1s not subdivided. A key to these species may be found in Harden & Rodd (1990).
40 Austrobaileya 5(1): 39-51 (1997)
Taxonomy
Key to native and naturalised taxa of Rubus in Australia. Naturalised taxa are marked with an asterisk (*)
1. Plants herbaceous, without prickles, carpels 6-10 ......00 0... cece eee R. gunnianus Plants woody, with prickles, carpels 25-600 .. 0... ee ee eens 2
2. Vines, without differentiated stems or canes; inflorescence axillary, race-mose; flowers UNISeEXUAal 1... ee eee ce ee ene tena eee ensneees 3 Sprawling shrubs, with stout sterile canes (primocanes) and flower-bearing side-shoots (floricanes); inflorescence terminal, HuiHCulate: TOWELS DISEMUAL 6 icivre.c ace a2ly 4 aber Tease acu dsk ao PaaS ea pelb dine SR we EM 4
3. Undersides of leaflets densely hairy throughout; margins with 5—7 teeth/cm; stipules present; carpels not glandular; aggregate fruits with TOS Osea CAPDIGS fon es a Wise ah TM eb slo Gack 0 Wty SiN wees Toieeteapert R. moorei Undersides of leaflets glabrous except for domatia and hairs along veins; margins with 3—5 teeth/cm; stipules absent; carpels glandular; aggregate
fruits with 35-65 glabrous carpidS ....... 0... ec eee eee R, nebulosus 4. Leaves on primocanes (and floricanes) simple... 1... cc ee teens 5 Leaves- on, priimniOcantes COMIPOUIL 5-55 FR cg bevrgiess 8 © ace nce eh gohe® scatewre Cs adnan ene anne 7
5. Stipules up to 15 x 15 mm, divided almost to their base, filamentous; leaves (especially young leaves) with a mixture of short and long hairs on
aa I SUET ACE oe cae etal a Nice ce 4 Sid ed eygivirete de, Vlorh Gowicers tebe ole Re ag Boa abel ac *R, alceifolius Stipules up to 12 x 6 mm, undivided at base, not filamentous; leaves with only long hairs on adaxial surface, or glabrous ......... 0. ccc eee ees 6
6. Most or all leaves prominently 3-lobed, with end lobe about two-thirds the length of the leaf; indumentum on petioles rather sparse, mostly appressed; petals mostly pink ..............000- R. moluccanus var. trilobus Leaves unlobed or lobes <1 cm deep, never prominently 3-lobed; indumentum on petioles dense, mostly spreading; petals white Suesartan ete Se talata: ara lanty a Beat ately leek gente desta ite wee erat R. moluccanus var. moluccanus
7. Leaves on primocanes pinnate or trifoliolate 0.0... eee ens 8 Leaves on primocanes pedate or palmate... 0. eee eee eee ye. 14
8. Leaflets 3—5, white or yellowish underneath, due to dense cover of short
hairs-obseuring the tearlet-surtace).... 4: cease beter tre oe mee Ob ek ag Eo ee 9 Leaflets S—9, green underneath, glabrous or hairy, surface of leaflets not ODSCUPEE OY AHS ose 4 ciate digesta ate eine a dh eke sh anlar Ween’ oo SMU: eH eee 1]
9. Branchlets with erect reddish hairs or bristles; petals white; fruits yellow ... *R. ellipticus Branchlets with appressed white hairs; petals pink or red; fruits (when TOTES CER NICER <a of Wes cid. ea + Reng endl Sane aA ae een renegades Bent anche nat Sorted oe BAe ae 10
10. Stipules 2-lobed, occasionally 3 or 4-lobed, fruits never formed............. R. x novus Slipules entines Huits LOM! 5.4 .yeycace dees ecd Quecerergew bes hu edeete Racer aie R. parvifolius
Bean, Rubus subg. Malachobatus 4] 11, Branchlets glabrous; leaflets glabrous or sparsely hairy on veins ...........0 00 cee. 12 Branchlets hairy; leaflets hairy throughout ......... 0. eee eee 13
12.Most carpels glandular; leaflets 14-27 mm wide; petals hairy on
both surfaces ......... 0.0000 ceeeeece
* «© © # # FF *® &#@ &#@ @© BF F © BB FF F&F ££ £ © 8 Ff
R. queenslandicus
No carpels glandular; leaflets 20-48 mm wide; petals glabrous on inner
MU LAC hs bd edhe die eat iters oe ain Lele dae
13. Flowers with 5 petals .......... 000. ee. Flowers with 9-13 petals ...............
14, Leaflets deeply incised or pinnatisect .....
= «© 8#® F FB @®© FB & *& & & & FF * & &£& &£& FF FF #& 8 &F§ FF FF FF BH FF FE
’' &+ & &§ * *# * 8 8 8 Ff
Leaflets often lobed, but never deeply incised or pinnatisect............ 0.0000 ee eee 15
15. Floricanes pruinose; mature fruits red ..... Floricanes not pruinose; mature fruits black
* ££ # © © #&* © © © © &© B&F # #@© @© @ & 8 8© #8 #@ @ © © b8 wp 8 # &£ © © © & & F 16
16. Lower surface of leaflets green and stems without glandular hairs; petals
ELSG mm iong: oo of. ep ae we tee eens
Pe meee at tert ee at ee *R. bellobatus
Lower surface of leaflets glaucous or stems with glandular hairs; petals
OSM AML IONG whe et Peele HORT RAD
Rubus subg. Malachobatus (Focke) Focke, Bibhoth. Bot. 72: 41 (1910); R. sect. Malachobatus Focke, Abh. Naturwiss. Vereine Bremen 4: 187 (1874). Type: not designated
About 127 species, mostly eastern and south- eastern Asia, also Malesia, Australia, western Pacific islands.
Shrubs, prickles present; leaves mostly simple, entire or lobed. Glands absent. Stipules free, on the twigs near the petiole base, mostly more or less divided. Inflorescence terminal, paniculate. Flowers mostly bisexual, carpels mostly 30-180; fruits red, succulent.
Rubus alceifolius Poir. in Lam., Encycl. 6: 247 (1804); R. moluccanus var. alceifolius (Poitr.) Kuntze, Meth. Sp.-Beschr. Rubus 56 (1879). Type: Java, 1766-69, P. Commerson s.n. (holo: P—JU, microfiche!).
For synonymy, see Kalkman (1984).
Illustrations: W.T.Parsons & E.G.Cuthbertson, Noxious Weeds of Austral. (1992: 576); D.L.Jones & B. Gray, Climbing PI. in Austral. (1988: 331).
sa GMAIL oS Bete ROG Si AT Ge *R, fruticosus agg.
Erect shrubs, sometimes semi-scandent, up to 6 m high. Stems hairy, with stout, curved to almost straight prickles to 4 mm long. Leaves simple, Petioles 35-55 mm long, terete, hairy, with a few stout prickles. Stipules rather caducous, orbicular in outline, to 15 mm long, 15 mm wide, divided almost to the base, with linear lobes; attached to stem at petiole base. Blades of leaves broadly ovate to orbicular, 85—150(-200) x 70—130(-200) mm, shallowly 5—7-lobed; upper surface green, hairy, comprising sparse long straight hairs and dense short stiff hairs; lower surface brown or yellowish, densely hairy, with short curly hairs almost or completely obscuring the leaf surface, and longer straight, spreading hairs on the veins; glands absent; base cordate; apex acute; margins serrate, with serrations 1—1.5 mm deep. Leaves palminerved at base, then penninerved with 2—3 pairs of lateral veins, terminating at a major tooth; venation reticulate, midrib and lateral veins flat to slightly impressed above, raised below, prickles occasionally present. Inflorescences terminal or in upper leaf axils, up to 100 mm long, each with up to LO flowers. Bracts broadly ovate in outline, to 17 mm long, deeply pinnatisect, segments linear, acute, hairy outside, glabrous inside. Pedicels terete, 5-14 mm long, sessile glands absent. Flowers
42
bisexual. Flower buds globular, hypanthia shallowly convex to almost flat, densely hairy, unarmed or with a few short prickles. Sepals persistent, ovate, 9-10 x 5.5—7 mm, outer ones with S—7 prominent linear lobes, inner ones not or scarcely lobed; sepals recurved at fruiting stage, and becoming somewhat larger; with dense white hairs on both surfaces, prickles absent. Petals not persistent, orbicular, shortly clawed, 7—9 x7—8.5 mm, glabrous, white, apex obtuse. Stamens 200-280; filaments up to 5.5 mm long, glabrous; anthers c. 0.75 mm long, versatile, dorsifixed, with several hairs at apex. Carpels 110-180, glabrous, glands absent; ovaries c. 1.0 mm long, styles c. 0.9 mm long. Carpels extending to base of torus, annulus of hairs present at base of torus. Collective fruit globular,c. 16 <x 16 mm when dried, c. 25 x 25 mm when fresh, exocarp red at maturity. Endocarpids reniform to ovoid, rugose, c. 2.5 x 1.7 x 1.3 mm.
Selected spectinens: Queensland. Cook Districr: 6 km E of Babinda, Dec 1992, Bean 5332 (BRI); c. 2 km SW of Bramston Beach on read from Bruce Hwy, Aug 1989, Blaxell 89/160 et al. (NSW); Palmerston Highway c. 10 km from Millaa Miilaa, Nov 1979, Clarkson 2731 (BRD; c. 40 km SE of Atherton at Josephine Falls, Aug 1981, Croat 52636 (NSW); c. 11 mls [18 km] from Innisfail neat Waugh’s Pocket, May 1952, Everist 5087 (BRD; S.F. 310 Gadgarra, track to Yeti Ridge, end of Fuller road, Jul 1995, Forster PIF17129 & Figg (BRI, K, MEL, NSW, QRS); Swipers flat, Wooroonooran N.P., Jul 1995, Forster PIFi7148 & Figg (BRI, MEL, NSW); Churchill Ck, Churchill LA, SF 143, Jul 1995, Forster PIF17199 & Figg (BRI, MEL, NSW, QRS); Cape Kimberley road, Nov 1995, Forster PIF 18178 (BRI, QRS); SFR 755, Bartle Frere, Boonjee L.A., Mar 1993, Gray 5642 (QRS); Danbulla, May 1961, Zyland 1859 (BRI); Cucantia, near junction of Russell and Mulgrave Rivers, Thorne 23132a et al. (BRI); Bellenden Ker, Mar 1922, White 1261 (BRI); Babinda, Jan 1923, White sn. (BRI). NortH KeNNepy DisTRICcT: i km W of Paluma, NW of Townsville, May 1992, Bean 4412 (BRI, K, MEL, NSW); Tam O’Shanter S.F. 1137, Mission Beach-Tully road, Jul 1995, Forster PIF17356 (BRI, MEL, NSW, QRS); 14 km NW of South Johnstone, Oct 1988, Jessup GJM2514 et al. (BRI); 11 km N of Tully on Bruce Highway, Apr 1975, Orchard 4657 (BRI); Mission Beach, Feb 1980, Stanfey 80255 (BRI). Port Curtis District: Quinns Gap road, c. 12.5 km NW of Carmila, Jul 1994, A¢eDonald 6056 et al. (BRD.
Distribution and habitat: R. alceifolius is indigenous to south-east Asia, from southern China to Indonesia, as far east as Sulawesi (Celebes). Naturalised populations in Australia extend from Cape Tribulation to Tully, with two disjunct populations further south at Paluma and Carmila (Map 1). It grows in
Austrobaileya 5(1): 39-51 (1997)
thickets on the edges of mesophyll or notophyll rainforest, and on other disturbed sites where rainforest once grew.
Phenology: Flowers have been recorded throughout the year; fruits have been recorded between May and August.
Notes: R. alceifolius is closely related to R. moluccanus, but can be readily distinguished by the deeply divided bracts and stipules with filiform segments, the leaves which are 5—7-lobed and often orbicular in outline, the globular flower buds, the indumentum on the adaxial leaf surface which comprises a mixture of long and short hairs (long hatrs only in R. moluccanus) and the greater number of carpels and stamens per flower.
I support the widely held belief that this species is naturalised in Australia. R. alceifolius was not recorded by F.M.Bailey during the Bellenden Ker Expedition in 1889, nor in Bailey (1900). It was first collected in Australia by C.T.White in March 1922, from Bellenden Ker (presumably near the railway siding of that name), and shortly after from Babinda. The species is apparently continuing to spread, as some recent collections are a long way from the first recorded locations.
R. alceifolius is considered a serious weed in Eacham, Johnstone and Mulgrave Shires, and is a Declared Plant, Category P3 under the Queensland Rural Lands Protection Act (1985-88), where ‘infestations must be reduced’,
Rubus moluccanus L., Sp. Pl. 1197 (1753). Type: plate 47, fig. 2, in Rumphtus, Herb. Amboin. 5: 88 (1747).
Rubus haskarlii subsp. dendrocharis Focke, Biblioth. Bot. 72: 99 (1910); R. dendrocharis (Focke) Focke, Bot. Jahrb. Syst. 54: 70 (1916);R. moluccanus var. dendrocharis (Focke) P.Royen, Phan. Monogr. 2: 106 (1969). Type: Bele River, 18 km NE of Lake Habbema, Nov 1938, Brass 11387 (neo: A, fide Royen (1969); iso: BRI).
For further synonymy, see Kalkman (1984).
Bean, Rubus subg. Malachobatus
Erect shrubs, sometimes semi-scandent, up to 6 m high. Stems hairy, with stout, curved to almost straight prickles to 2 mm long. Leaves simple. Petioles 22-52 mm long, slightly grooved on upper surface, hairy, with a few stout prickles. Stipules caducous, elliptic in outline, to 12 mm long, 6 mm wide, undivided at base, with linear lobes; attached to stem at petiole base. Blades of leaves ovate to broadly ovate, 45-150 <x 40-125 mm, shallowly 3—S-lobed or prominently 3-lobed, or sometimes entire; upper surface green, sparsely hairy, comprising long straight hairs only; lower surface brown or yellowish, densely hairy, with short curly hairs obscuring the leaf surface, and longer straight, appressed or spreading hairs on the veins, glands absent; base cordate to auriculate, but adjacent lobes only rarely touching each other; apex acute or occasionally obtuse; margins serrate, with serrations 1-2 mm deep. Leaves palminerved at base, then
43
terminating at a major tooth; venation reticulate, midrib and lateral veins flat to slightly impressed above, raised below, prickles occasionally present. Inflorescence terminal or in upper leaf axils, up to 120 mm long, with up to 15 flowers. Bracts ovate to broadly ovate in outline, to 9 mm long, deeply pinnatisect, segments linear, acute, hairy outside, glabrous inside. Pedicels terete, 6-11 mm long, sessile glands absent. Flowers bisexual. Flower buds ovoid, hypanthia shallowly convex to almost flat, densely hairy, unarmed. Sepals persistent, deltate, 6-9 x 4-5 mm, without an acumen, entire or with 3—5S short lobes, recurved at fruiting stage, and becoming somewhat larger; with dense white hairs on both surfaces, prickles absent. Petals not persistent, broadly ovate, not or shortly clawed, 6—8 x 4—5.5 mm, glabrous or sparsely hairy, white or pink, apex obtuse. Stamens 60—110; filaments up to 4mm long, glabrous; anthers c. 0.75 mm long,
penninerved with 5—7 pairs of lateral veins, — versatile, dorsitixed, glabrous or with several
hairs at apex. Carpels 35—70, glabrous, glands absent; ovaries c. 0.9 mm long, styles
ST aeenesee MDAOLT Wseueeacesesceneeer gsersfuoeee! ne. | Net CC _ Pid a SR aR EREEERSCEED BEER GEC CEE EE EEE EEE TERRE
ssdastcnee fat serataceel
155 ! 10 l |
SEESEGGR SRE a PREECE 3 | EEC CCC ECC
ce SECC | sth
sarees ECE eee ce \ LN | une 2 ! = -EEEEE Ra hs
SL yy TE ed PEE rh
y
=euanenue CCC ies , Ss FECRECECE EEE SPEREE ECE pe
Map 1. Distribution of A Rubus alceifolius and
kK | COE +N PPR rare EEE ) gannEre SeERGP/) Pere Ha
seruentorec tout batanfhr7/safoasastacter gat
: | } 35
Ha
= t- i - —s
ae R. Bun fi | Gnhnus,
44
c. 7.5 mm long. Carpels extending to base of torus, annulus of hairs present at base of torus. Collective fruit globular or oblate, c. 10 x13 mm when dried, c. 12 x 16 mm when fresh, exocarp red at maturity. Endocarpids reniform to ovoid, lacunose, 6.2.3 * 1.5 x 1.5 mm.
Two varieties occur in Australia; R. moluccanus var. moluccanus and R. moluccanus var. trilobus.
Rubus moluccanus L. var. moluccanus
Rubus hillii F.Muell., Trans. & Proc. Philos. Inst. Victoria 2: 67 (1857). Type: Queensland. Moreton District: Brisbane River, post Feb 1855, W. Hill s.n. (holo: MEL [MEL 31307]; iso: K).
Rubus capricorni Focke, Annuaire Conserv. Jard. Bot. Geneve 20: 106 (1917). Type: Queensland. Cook District: Kamerunga near Cairns, 1885-89, O. Warburg 18628 (holo: B, destroyed).
Rubus moluccanus var. austropacificus P.Royen, Phan. Monogr. 2: 113 (1969) (‘austropacifica’). Type: Bougainville. Tuareruku village W of Toiumonapu Plantation, S of Kieta, 4 July 1963,P. van Royen NGF 16444 (holo: L n.v.; iso: BRI).
Leaves 85-130 x 60—125 mm, shallowly 3—5-lobed, with sinuses <]0 mm deep, or entire; young stems, petioles and veins of the lower leaf surface with erect or spreading, long straight hairs, as well as short curly hairs which obscure the lamina; petals white.
Selected specimens: Queensland. Coox District: c. 14 mls [23 km] NNW of Daintree, Nov 1967, Boyland & Gillieatt 415 (BRI, MEL, NSW); Black Mountain road, near Kuranda, Jan 1968, Brass 33753 (QRS); c. 1 km S of Kuranda on road to Wrights Lookout, Jan 1980, Clarkson 2752 (BRI, QRS); near Mt Haig, c. 4.5 km NNW of Danbulla, Nov 1979, Clarkson 2722 (BRI, L, QRS); 32 km along road to Leo Creek, Mcliwraith Range, Jun 1995, Forster PIF16821 (BRI, K, MEL, NSW, QRS); Home Rule, Jul 1995, Forster PIF 17289 & Figg (BRI, MEL, NSW, QRS); Nissen Creek, Mt Perseverance road, Jul 1995, Forster PIF17178 & Figg (BRI, QRS); Timber Reserve 14, Kesteven, Oct 1981, Hyland 11152 (QRS); Noel L.A., N of Tinaroo Falls Dam, Nov 1981, Kanis 2152 (BRI, CANB). Norry Kennepy District: just W of Paluma, NW of Townsville, Feb 1992, Bean 3948 (BRI);
Austrobaileya 5(1): 39-51 (1997)
top of Seaview Range escarpment, Trebonne-Mt Fox road, Apr 1985, Rodd 4485 & Hardie (BRI, MEL, NSW); headwaters of Dryander Creek, Mt Dryander, Feb 1971, Webb & Tracey 10073 (BRI, CANB); Wilkin Hill, Hinchinbrook Is., May 1972, Webb & Tracey 12011 (BRI, CANB, QRS). SouTH KENNEDY District: Mt Blackwood, Mar 1987, Thompson 79 (BRI). Wipe Bay DIstTRIct: Gheerulla Falls, Mapleton S.F., Nov 1990, Bean 2684 (BRI, NSW); Kin Kin, Jan 1917, White s.n. (BRI, K). Moreton District: Mt Cougal N.P., Upper Currumbin Creek, Mar 1997, Bean 11765 (BRI, MEL, NSW); 2.5 mls [4.0 km] SW of Nambour, Jan 1970, Parsons s.n. (BRI); Mt Glorious, Dec 1950, StJohn 24402 (K). New South Wales. Nortu Coast: Richmond River, undated, coll. unknown (NSW [NSW394747]); Clarence River, undated, 7Adoore (MEL [MEL 31326]).
Distribution and habitat: R. moluccanus var. moluccanus 1s native to south-east Asia, including Thailand, throughout Malesia and extending to Fiji (Salkman 1984). In Australia, it is indigenous to coastal areas from northern Queensland to northern New South Wales (Map 2), although no collections have been made in New South Wales for over 100 years. It grows in open locations within or fringing notophyll or mesophyll rainforest, or sometimes in open forest with (for example) Eucalyptus grandis W.Hill ex Maiden at altitudes from sea-level to 1140 metres.
Phenology: Flowers and fruits have been recorded for most months of the year.
Note: Bentham (1864) recorded R. moluccanus for the Northern Territory based on a specimen with the label ‘Port Efsington |[Essington?], Armstrong’. This distributional record was subsequently repeated by Ewart & Davies (1917) and appears in several popular publications of recent years. However, the record appears to be erroneous. Firstly, the specimen in question belongs inR. moluccanus var. discolor (BI.) Kalkman, a taxon not recorded for Australia, but common in Malesia and the islands of the western Pacific. Secondly, R. moluccanus is not otherwise known from the Northern Territory, based on both herbarium records and the comprehensive atlas provided by Liddle et al. (1994). Furthermore, no Rubus species has been seen by botanists of the Northern Territory Herbarium (DNA) (C. Dunlop, pers. comm.).
my SL Mec eye emp ee Md ae WE NT a
Bean, Rubus subg. Afalachobatus 45
Armstrong is known to have collected in foliis conspicue trilobis lobo suo terminali Timor for some years (Britten & Boulger 1931), longitudine dimidium folii excedenti, and it is likely that the‘Port Essington’ specimen indumento plerumque adpresso et petalis has originated from Timor, and that a saepe roseis, differens. Typus: New misplacement of specimen labels has occurred. South Wales. NoRTHERN J'ABLELANDS:
Giro State Forest, between Walcha and Typification: The holotype of R. capricorniwas Gloucester, 10 December 1996, destroyed during World War II. Isotypes were A.R. Bean 11477 (holo: BRI; iso: K, L, sought from E, K, BM, FI, G and W, but without MEL, NSW).
success. The protologue is sufficiently detailed
to allow placement of this name as a synonym [R. Ailfii auct. non F.Muell.|
of R. moluccanus var. moluccanus. Illustration: G.R. Cochrane et al., Fl. & Plants
Conservation status: R. moluccanus var. of Victoria & Tasmania, p. 97 (1980), as
moluccanus is a widespread and common — &. Aillii.
taxon. No conservation coding is
recommended. Leaves 45-110 x 40-100 mm, deeply 3-lobed, with sinuses >10 mm deep, and terminal lobe
R. moluccanus var. trilobus A.R.Bean var. —c. two-thirds of leaf length; young stems,
nov. varietati moluccanae affinis, sed petioles and veins of the lower leaf surface with
appressed or sometimes spreading, long straight hairs, as well as short curly hairs which obscure
SEE the lamina; petals usually pale or deep pink, but sometimes white. |
Map 2, Distribution of Rubus moluccanus var. moluccanus.
46
Selected specimens: Queensland. Cook District: Sylvia Creek c. 4 km W of Atherton, Nov 1979, Clarkson 2742 (BRI, QRS), NortH KENNEDY District: Mt Graham, Rockinghams Bay, Dallachy s.n. (MEL); Magnetic Island, around summit of Mt Cook, Aug 1982, Sandercoe 903 (BRI). SoutH Kennepy District: Carlisle Island, 35 km N of Mackay, Sep 1986, Sharpe 4466 et al. (BRI). Porr Curtis District: Mt Maria, c. 65 km NW of Bundaberg, Novy 1993, Bean 7003 (BRI, DNA); Rockhampton, Apr 1868, O’Shanesy 99 (MEL). Wipe Bay District: Middle road, near Kenilworth-Jimna road, Dec 1990, Bean 2710 (BRI, CANB, MEL). DarumNc Downs District: Spicers Gap, Main Range N.P., via Aratula, May 1995, Bean 8643 (BRI, MEL). Moreton District: Mount Mistake, Nov 1930, Hubbard 5217 (BRI, K). New South Wales. NorTHERN TABLELANDS: Mt Boss SF, 45 km NW of Wauchope, May 1975, Coveny 6375 et al. GS). NortH Coast: Dalmortan State Forest, SW of Grafton, Aug 1994, Bean 7766 (BRI); Toonumbar, NW of Kyogle, May 1949, Constable NSW10543 (K, NSW). CENTRAL TABLELANDS: Murdering Gully, Kanangra, 10 miles [16 km] SE of Jenolan Caves, May 1965, Constable 5906 (K, NSW); foot of Katoomba Falls, Blue Mts, Dec 1950, StJohn 24604 (K, NSW). CENTRAL Coast: Bulli Pass, Apr 1896, Morrison 5271 CK, NSW). Sours Coast: Mt Dromedary, Courta Creek, Mar 1980, Craven 5422 (BRI, CANB, MEL, NSW); McCarthys Ck, Tantawangalo S.F., Dec 1990, Crawford 1249 (MEL, NSW). Victoria. Howe Ranges, N of Marshmead, Nov 1969, Beauglehole 31453 & Willis
Austrobaileya 5(1): 39-51 (1997)
(MEL); Scout Camp Rd, Major Ck crossing, 5.1 km N of Orbost, Aug 1979, Forbes 140 (MEL, NSW); Newtons Ck, near Cabbage Tree Creek, Dec 1910, St/o/in s.n. CK, MEL).
Distribution and habitat: R. moluccanus var. trilobus is endemic to Australia, and it extends from the Atherton Tableland in north Queensland to far eastern Victoria (Map 3). It commonly grows in gullies or sheltered situations in open eucalypt forest, and also on the edges of notophyll rainforest. In north Queensland it is confined to altitudes above c. 500 metres, but elsewhere it may extend to sea-level.
Phenology: Flowers and fruits have been recorded from every month of the year.
Notes: The rank of variety has been chosen for this taxon, because it differs from R, moluccanus sens. str. only in the leaf shape, characters of the indumentum and the often pink petals; and because this rank 1s compatible with the treatment of Kalkman (1984). The
113 120 125 (} 135 140
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Map 3. Distribution of Rubus moluccanus var. trilobus
135
140 Id5
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Bean, Rubus subg. Adalachobatus
name R. hillii has long been misapplied to this taxon.
Conservation status: R. moluccanus var. trilobus 1s a widespread and common taxon. No conservation coding is recommended.
Rubus x novus Kuntze, Meth. Sp.-Beschr. Rubus (1879). Type: New South Wales. NortH Coast: Hastings River, in 1862,H. Beckler s.n. (ecto, here designated: K).
Rubusx illegitimus Focke, Annuaire Conserv. Jard. Bot. Geneve 20: 106 (1917). Type: Queensland. DARLING Downs DistRIcT: Toowoomba, 1885-89, O. Warburg 18629 (holo: B, destroyed),
Rubus x novae-cambriae Gand., Bull. Soc. Bot. France 65: 25 (1918). Type: New South Wales. CENTRAL Coast: Menangle, Oct 1893, R.T. Baker s.n. (holo: LY nv., photo at NSW).
Sprawling shrubs, sometimes semi-prostrate, up to | m high. Stems hairy, with stout, curved to almost straight prickles to 2.5 mm long. Leaves up to 105 mm long, usually trifoliolate, rarely simple but then deeply 3-lobed. Petioles 10-44 mm long, grooved on upper surface, hairy, with a few stout prickles. Petiolules of lateral leaflets absent or up to 1 mm long, erooved, hairy; petiolules of terminal leaflets 0—11 mm long, grooved above, hairy. Stipules persistent, linear, to 12 mm long, 1.5 mm wide, usually 2-lobed, sometimes these lobes further divided; apex acute, attached at junction of petiole and stem. Blades of leaflets ovate, 25~75 X 2-50 mm, the terminal leaflet often larger than the lateral ones; upper surface green, sparsely hairy, comprising long straight hairs only; lower surface white or yellowish, densely hairy, with short curly hairs obscuring the leaf surface, and longer straight, appressed or spreading hairs on the veins; glands absent; base broadly cuneate or obtuse, sometimes oblique in lateral leaflets; apex acute; margins serrate, with serrations 1-3 mm deep. Leaflets penninerved with 4—6 pairs of lateral veins, terminating at a major tooth; venation reticulate, midrib and lateral veins impressed above, raised below, prickles occasionally present.
47
Inflorescence terminal or in upper leaf axils, up to 80 mm long, with up to 10 flowers. Bracts 3—5—partite, to 9 mm long, segments linear, acute, hairy outside, glabrous inside. Pedicels terete, 4-20 mm long, hairy, glands absent. Flowers bisexual. Flower buds ovoid, hypanthia shallowly convex to almost flat, hairy, unarmed. Sepals persistent, deltate, 5.5—7 x 2.5-3.5 mm, including a 0.5—2.5 mm long acumen, entire or occasionally with small lobes, recurved after anthesis; with dense white hairs on both surfaces, prickles absent. Petals not persistent, elliptical, clawed, 6-8 x 4—5 mm, glabrous or with some hairs on outer surface, pink, Stamens 50-80, glabrous; filaments up to 4.5 mm long, anthers c. 0.75 mm long. Carpels 40-50, glabrous, glands absent; ovaries c. 1.0 mm long, styles c. 5.5 mm long. Carpels extending to base of torus, annulus of hairs present at base of torus. Fruits never formed.
Selected spectinens: Queensland. SouTH KENNEDY District: headwaters of East Funnel Creek, SSE of Sarina, Sep 1993, Champion 862 (BRI). Port Curtis District: Reedy Creek, 8 km SE of Lowmead on Bundaberg road, May 1976, Rodd 3046 & Jacobs (NSW). Burnett DIstrict: Mt Perry, undated, Keys s.n. (BRI). Dartinc Downs District: Cherry Plain Lookout track, Bunya Mountains N.P., May 1996, Grimshaw PG2470 & Grimshaw (BRI, NSW). Moreton District: 6.5 km WNW of Bellthorpe Hall, towards Forestry station, Feb 1989, Bean 1004 (BRD; foothills of Mt Ballow, downstream from Grace’s Hut, Dec 1989, Bean 1287 (BRI, L, NSW); 7km NW of Mt Glorious, Jan 1990, Bean 1331 (BRI, L, NS W); Tamborine Mtn, Dec 1926, White 3344 (BRI). New South Wales. Nortu Coast: Maclean, Feb 1913, coll. unknown (NSW); Gloucester, Jan 1882, Betche s.n. (NSW); Ramornie, 3 miles [5 km] NW of Copmanhurst, Jul 1922, Blakely & Shiress s.n. (NSW); Laurieton (Camden Haven), Nov 1944, McComish s.n. (NSW). CENTRAL WESTERN SLOPES: Murrurundi, May 1986, Cherry s.n. (NSW), CENTRAL TABLELANDS: Bundancon, Feb 1897, Stee/ sn. (NSW). Centra Coast Pulbah Island, Lake Macquarie, Nov 1929, Chee/ s.n. (NSW); Bulli, Dec 1888, Fletcher s.n. (NSW); Point Clare, Jun 1897, Forsyth s.n. (NSW); Machin’s Crater, 5.5 miles [8.8 km] SSE of Glenbrook, Mar 1970, Pickard 953 (NSW), SoutH Coast. Kiah (or Towamba) River c. 2,5 km NE of Kiah, Nov 1976, Jacobs 2880 (NSW). Victoria. 4 km NE Goongerah, Orbost region, Jan 1986, Chesterfield 651 (MEL),
Distribution and habitat: R. x novus has been recorded along the eastern Australian coast, from Mackay in Queensland to eastern Victoria (Map 4). It grows close to notophyl! rainforest or in tall eucalypt forest, dominated by species such as Eucalyptus saligna Sm. and i. microcorys F,Muell.
48
Phenology: Flowers have been recorded between June and March.
Conservation status: R. X novus 1s a widespread and common taxon. No conservation coding is recommended.
Notes: R. x novus 1s a naturally occurring hybrid, between R. moluccanus var. trilobus (of R. subg, Malachobatus) and R. parvifolius L. (of R. subg. Idaeobatus). It is the only known hybrid between indigenous Australian Rubus taxa, and is perhaps the only naturally occurring inter-subgeneric hybrid for the genus Rubus, although Kalkman (1984) speculated on the existance of an inter-subgeneric hybrid for a specimen he collected (No. 4856). I have not seen this specimen.
Austrobaileya 5(1): 39-51 (1997)
R. x novus 1s sterile. While it flowers well, no fruits have ever been collected or reported, and I have never seen any, despite actively searching for them.
Typification: Kuntze (1879) clearly states that he saw a specimen from Hastings River, New South Wales at the Herbarium of the Royal Botanic Gardens, Kew and as the protologue agrees with this specimen, it is chosen as the lectotype for the name Rubus x novus.
The holotype of R. x i/legitimus was destroyed during World War II. Isotypes were sought by the present author from E, K, BM, FI, G and W, but without success. The protologue is sufficiently detailed to allow placement of this name as a synonym of R, X HOVUS.
Rubus subg. Diemenicus A.R.Bean subg, nov.
Map 4. Distribution of Rubus x novus
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“SESS MEPs Midiehelaeseame cena vea renieoenvea ve ene ge ore eerie corcenaaeer Sue EN
Bean, Rubus subg. Mdalachobatus
Herba rhizomate repente, sine aculeis; folia omnia radicalia, petioli e basi vaginantes, stipulae nullae; flores solitari1, unisexuales, carpellis 6 —10; glandulae breviter pedunculares in petiolis foliis pedunculis ovariisque adsunt; fructus rubri, succulent.
Typus: R. gunnianus Hook. A monotypic subgenus.
Rhizomatous creeping herb, prickles absent, leaves all radical, petioles sheathing at base, stipules absent; flowers solitary, unisexual, carpels 6 —10; shortly stalked glands present on petioles, leaves, peduncles and ovaries; fruits red, succulent.
Etymology: Named for Van Diemen’s Land, now known as Tasmania, where the type species is endemic.
Rubus gunnianus Hook., Icon. Pl. 3: t. 291 (1840). Type: Tasmania. Chilton, Surrey Hills, 16 February 1837, R. Gunn 271/1837 (lecto: K, here designated; isolecto: NSW [NSW385192]).
Illustrations: Cochrane et al., Fl. and Plants of Victoria & Tasmania, p. 112 (1980); Macoboy et al. (eds), Gard. Encyc. of Pl. & Flowers, p. 321 (1992),
Prostrate herbs, lacking prickles, rhizomatous. Vertical stems 1-10 mm long, rarely to 50 mm, glabrous, prickles absent. Leaves borne in clusters at each node; up to 60 mm long, mostly with 3 leaflets, but with unifoliolate leaves often present at base of cluster. Petioles 12-36 mm long, grooved on upper surface, sheathing at base with sparse, simple white hairs and stipitate red glands. Petiolules of lateral leaflets 0—-0.5 mm long, grooved, glands present; petiolules of terminal leaflets 0-3 mm long, grooved above, glabrous, glands present. Stipules absent. Blades of terminal leaflets broadly ovate, 9-19 x 7-14 mm, the lateral leaflets broadly ovate, 6-8 x 4.5—5 mm; glabrous or with sparse hairs along midrib, glands stipitate, red, turning black with age, rather sparse; base broadly cuneate or obtuse, sometimes oblique in lateral leaflets; apex acute; margins serrate to deeply lobed. Leaflets penninerved with 2—4 pairs of lateral veins,
49
branching before the margin and terminating at a sinus; venation reticulate, midrib and lateral veins impressed above, raised below. Inflorescence axillary, solitary. Bracts ovate, entire, to 6 mm long, glabrous or with sparse hairs and glands along margins, apex obtuse. Peduncies terete, 4-16 mm long, with white hairs, stipitate glands present. Flowers functionally unisexual. Flower buds globular, hypanthia obconical, sparsely hairy. Sepals persistent, deltate to ovate, 4—5.5 x 2.5—3.5 mm, entire; with sparse white hairs on both surfaces. Petals not persistent, narrowly elliptical, not clawed, 6-10 x 2-4.5 mm, glabrous, white. stamens 16-23, glabrous; filaments up to 2.5 mm long, anthers c. 0.75 mm long. Carpels 6—10, glabrous; glands numerous per carpel, shortly stalked; ovaries c. 0.8 mm long, styles c. 0.75 mm long. Torus small, hairy between the carpels; annulus of hairs absent at base of torus. Collective fruit globular, c. 10 x 10 mm when dried, of unknown size when fresh, exocarp red at maturity; carpids 2-4 per fruit. Endocarpids ellipsoidal, lacunose, c. 4.5 x 2.5 x 2.5 mm.
Selected specimens: Tasmania, NortH West: Waratah, Dec 1914, Lucas s.n. (NSW). Ringwood, Nov 1841, Milligan 152 (HO). BEN Lomonpb: summit of Mt Maurice, Dec 1979, Buchanan 71 CHO); Mt Barrow, Jan 1922, Rupp s.n. (MEL). West Coast: Trig station of Raglan Range, Dec 1984, Collier 156 (HO); Linda track, Jan 1903, Rodway 214 (HO); Lake Rhona, Denison Range, Jan 1977, Tyler sn. (HO), CENTRAL HicHLANps: Cradle Valley, Dec 1931, Anderson 9 CHO); Drys Bluff, NE face, just below the summit, Dec 1984, Buchanan 4932 (HO); below Eldon Bluff, Feb 1987, Buchanan 9970 CHO); Micklethwaite Marsh, Mar 1985, Morris 8535 (HO); King William Range, Nov 1933, Rodway 154 (HO), Mr. Fie_p: Lake Fenton, Nov 1914, Gibbs 6508 (BM); Mt Field N.P., Jun 1979, Noble 28843 (HO); near Lake Dobson huts, Mt Field N.P., Jan 1978, Smith 237 (HO). Mr. WELLINGTON: Mt Wellington, Jan 1892, Bailey & Shirley (BRD; Mt Wellington, Feb 1891, Rodway s.n. (MEL). SouTu WEst: Southern Jubilee Range, Jan 1985, Buchanan 5396 (HO); 3 km ESE of Mt Mueller, Dec 1990, Collier 4958 (HO); Hartz Mountain, Apr 1901, Rodway 214 (HO).
Distribution and habitat. Rubus gunnianus 1s endemic to the island of Tasmania, where it is apparently common in many places between altitudes of 620-1280 metres (Map 1). Its most commonly recorded habitat is amongst alpine or sub-alpine shrubbery, but it is also recorded from Buttongrass (Gymnoschoenus sphaerocephalus (R.Br.)
50
Hook.f.) plains, shady gullies and dolerite cliffs.
Phenology: R. gunnianus produces flowers from November-February and fruits from January-April.
Notes: R. gunnianus has no close relatives. While Focke (1910) placed it in R. subg. Dalibarda, he also stated that “R. gunnianus and R. dalibarda truly constitute two characteristic subgenera”.
A new subgenus is here erected to accommodate R. gsunnianus, as tt has some characteristics unique for Rubus, or at least its non-woody members. These include 1) leaves in rosettes, often arising directly from the rhizome, or with short erect branches; 11) absence of stipules (unlike all other creeping Rubus); 111), broad petioles sheathing at their bases; Iv) prominent glands on the carpels, unlike all other herbaceous species of Rubus; v) flowers often (perhaps always) unisexual, with either stamens or carpels being imperfectly developed (Curtis 1956; Rodway in litt.), and this can be observed in the herbarium. All other herbaceous Rubi are bisexual, except R. chamaemorus L., which is dioecious (Bailey 1941); vi) The major lateral veins 1n the leaves terminate at a sinus. This occurs in no other Rubus species that I have examined, and hence this is possibly another unique characteristic, although the available data are very limited.
According to Focke (1910),2. gumnianus is the smallest of all Rubus species.
Typification: The lectotype comprises four separate pieces (two bearing flowers and two bearing fruit), in a horizontal row across the centre of the sheet, and labelled “Mr Gunn 271/ 1837, Van D. Land, ripe fruit at Chilton, Surrey Hills, 16/2/37”. Two of these pieces appear to form the basis of the illustrations accompanying the protologue. Other specimens mounted on the same sheet are dated 1840 and 1845, and could not have been used by Hooker in drawing up his description. Hence they are excluded from forming part of the type.
The isolectotype at NSW consists of the mounted material only. The material in the
Austrobatleya 5(1): 39-51 (1997)
packet was collected by J. Milligan and 1s hence excluded.
Conservation status: R. gunnianus is reported to be common throughout its habitat, and 1s present in at least two National Parks. No conservation coding is recommended.
Acknowledgements
Specimens from several herbaria have been examined; I would like to thank the Directors of AD, BM, HO, K, MEL, NSW, PERTH and QRS for specimen loans. Laurie Jessup and Barry Conn searched for type specimens on my behalf during their terms as Australian Botanical Liaison Officer. Philip Sharpe kindly translated Kuntze’s paper relating to R. x novus. Paul Forster collected specimens from north Queensland at my request. Gordon Guymer commented on a draft of this paper. Peter Bostock assisted in the translation of some Latin text, and provided the Latin diagnoses.
References
BalLtey, F.M. (1900). Rubus. In The Queensland Flora 2: 526-8. Brisbane: H.J. Diddams & Co.
Bamey, L.H. (1941). Species Batorum: The Genus Rubus in North America (north of Mexico). Gentes Herbarum 5: 1-932.
Bean, A.R. (1995), A revision of Rubus subg. Micranthobatus (Focke) Kalkman (Rosaceae) in Australia. Austrobaileya 4; 321-8.
—— (1997), A Revision of Rubus subg. Idaeobatus (Focke) Focke (Rosaceae) in Australia. Austrobaileya 4: 677-89.
BENNETT, E.M. (1987). Rosaceae. In Marchant, N.G. et al., Flora of the Perth Region, 1:207—11. Perth: Western Australian Herbarium.
BENTHAM, G. (1864). Rubus. In Flora Australiensis 2: 429-32, London: L. Reeve & Co.
Britten, J. & Boutcer, G.S. (1931). A Biographical Index of Deceased British and Irish Botanists, 2nd edition. London: Taylor & Francis.
Curtis, W.M. (1956). The Student’s Flora of Tasmania, Part 1.Tasmania: Government Printer.
Ewart, A.J. & Davies, O.B. (1917). The Flora of the Northern Territory, Melbourne: Minister for Home and Territories.
ee to eA SE RE EM a i i
Bean, Rubus subg. Malachobatus
Focke, W.O. (1910), Species Ruborum Monographiae generis Rubi Prodromus. Bibliotheca Botanica 17 (72 Part 1): 1-120.
GLEASON, H.A. (1952). New Britton & Brown Illustrated Flora 2: 318. New York: New York Botanical Gardens
HARDEN, G.J. & Ropp, A.N. (1990). Rosaceae. in G.J, Harden (ed.), Flora of New South Wales 1; 528-50. Sydney: New South Wales University Press.
KALKMAN, C. (1984), The Genus Rubus (Rosaceae) in Malesia 2. The subgenus Ma@lachobatus. Blumea 29: 319-86.
Kuntze, O. (1879), Methodik der Speciesbeschreibung und Rubus. Leipzig: Arthur Felix.
Lippe, D.T., RUSSELL-SMITH, J., Brock, J., LeEAcH, GJ. & Connors, G.T. (1994). Atlas of the Vascular Rainforest Plants of the Northern Territory. Canberra: Australian Biological Resources Study.
Royen, P. vAN (1969). The Genus Rubus (Rosaceae) in New Guinea. Phanerogamarum Monographiae Tomus II. Lehre: J. Cramer.
Scoacan, H.J. (1978). Rosaceae. in The Flora of Canada 3: 898-967, Ottawa: National Museums of Canada.
Symon, D.E. (1986). Rubus, In Jessop & Toelken (eds.) Flora of South Australia 1: 446-9.
THomeson, M.M. (1997). Survey of Chromosome Numbers in Rubus (Rosaceae: Rosoideae). Annals of the Missouri Botanic Garden 84: 128-64.
Wess, C.J., SYKES, W.R. & GARNOCK-JoNEs P.J, (1988). Rubus, in Flora of New Zealand, Volume IV, Naturalised Pteridophytes, Gymnosperms, Dicotyledons. Christchurch: Botany Division DSIR.
51
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Madangia inflata (Asclepiadaceae: Marsdenieae), a new genus and species from Papua New Guinea
Paul I. Forster, David J. Liddle and Iris M. Liddle
Summary
Forster, Paul f., Liddle, David J., Liddle, Iris M. (1997). Madangia inflata (Asclepiadaceae: Marsdenieae), a new genus and species from Papua New Guinea. Austrobaileya 5(1): 53-57. The new genus Madangia is described with the sole species AZ inflata P.1.Forst., D.J.Liddle & LM.Liddle. M. inflata 1s thus far known only from the Madang Province of Papua New Guinea. The genus is
closely allied to Hoya R.Br. to which it is compared.
Keywords: Asclepiadaceae, Marsdenieae, Hoya, Madangia - Papua New Guinea; Madangia inflata.
Paul I.Forster, Queensland Herbarium, Meiers Road, Indooroopilly, Queensland 4068, Australia David J.Liddle, Iris M.Liddle, P.O, Box 794, Mareeba, Queensland 4880, Australia
Introduction
The authors are working towards a revision of the genera Hoya R.Br. and Dischidia R.Br. (both Asclepiadaceae: Marsdenieae) as they occur in Australia and Papuasia. In 1992 the first two authors collected numerous live material of Hoya in north-east Papua New Guinea, particularly in Madang Province, One of the plants, collected at the time without flowers and with the vegetative appearance of a species of Hoya, has now subsequently flowered in cultivation and is not referable to that genus. This plant represents not only an undescribed species, but an undescribed genus, which although allied to Hoya, differs in numerous ways from all taxa included in that genus. This species is described as Madangia inflata in the present paper and its affinities discussed. There appear to be no previous herbarium collections of this plant that has rather remarkable, showy, white, globose flowers and is of considerable horticultural merit.
Madangia P.I.Forst., D.J.Liddle et LM.Liddle genus novum. Genus singulare in Asclepiadaceis: Marsdenieis per corollam globosam et formam loborum coronae staminalis. Lobi coronae staminalis membranei et sibi contigui (autem non
Accepted for publication 16 July 1997
connati) secus longitudinem paene omnino praeter apices supernos lanceolatos prope apicem styli et margines infernos columna staminali multo longiores. Margo infernus quuisque columnae staminalis valde recurvus in extremitate necnon segmenta membranacea involutiones corollae interposita format.
A genus unique within the Asclepiadaceae: Marsdenieae by virtue of the globose corolla, and the form of the staminal corona lobes. The staminal corona lobes are membranous and contiguous (but not fused) to each other for almost their entire length, apart from the upper lanceolate apices near the style head and the lower edges that are much longer than the staminal column. The lower edge of each staminal corona lobe is strongly recurved at the end and forms a membranous frill positioned between involutions of the corolla wall.
Type species: Madangia inflata P.1.Forst., D.J.Liddle & I.M.Liddle
Epiphytic twiner; latex white. Roots fibrous. Leaves petiolate; lamina fleshy, hairy; with colleters. Inflorescence deciduous, usually single at a node, umbelliform to racemiform, positively geotropic. Flowers pedicellate. Sepals 5. Corolla globose, fleshy-membranous;
54
tube much inflated, of 5 completely fused petals; lobes free at tips, valvate. Annular and corolline coronas absent; staminal corona of 5 membranous lobes that are contiguous with but not fused to each other, each lobe attached to the back of each anther that make up the fused staminal column, lower edge of lobe strongly flanged away from central part of flower and strongly recurved with a somewhat frilled margin that is positioned between involutions of the corolla wall; interstaminal corona absent. Stamens connate into gynostegium (staminal column), not markedly capitate. Anthers 2-locular with terminal appendage; pollen in linear tetrads; pollinaria with 2 erect, oblong pollinia with a pellucid germinating mouth on outer margin; corpsculum oblong-ovate; caudicles not winged, not geniculate. Style-head rounded. Follicles and seeds not seen.
A monotypic genus, endemic to Papua New Guinea.
Etymology: The generic name is formed from the geographical region of Madang Province in north-east Papua New Guinea.
Madangia inflata P.I.Forst., D.J.Liddle et I.M.Liddle sp. nev. Volubilis lenta; indumento trichomatum incoloratum simplictum multicellularum. Caules usque ad metra pluria longi ad interque internodos radicantes, trichomatibus sparsis; internodia usque ad 200 mm longa et 2 mm diametro, lenticellata demum. Petioli cylindrici, 8-12 mm longi, 1.5-3.5 mm diametro, leviter sulcati supra, trichomatibus sparsis antroris. Colletri rotundati 1 vel 2 basi laminae. Lamina folii elliptica usque ad elliptico-ovata, succulenta, usque ad 120 mm longa et 60 mm lata, apice acuta usque ad breve acuminata, basi cordata lobis superpositis; venatio ex venis 6 vel 7 lateralibus arenatis et et venis interlateralibus reticulatis mndistinctis constans; pagina supera viridis venatione + obscura trichomatibus dispersis; pagina infera pallide viridis venatione indistincta trichomatibus dispersus usque ad sparsis. Pedunculi florentes 8-22 mm longi,
Austrobaileya 5(1): 53-57 (1997)
1.5—2 mm diametro, trichomatibus dispersis floribus usque ad 9 ornati; bractea ovato-truncata, 0.4-0.5 mm longa, 0.5-0.8 mm lata, sparse ciliata. Flores 12-14 mm longi, 17—18 mm diametro; pedicelli 32-45 mm longi, 1.4-1.5 mm diametro, glabri. Sepala triangularia, c. 1 mm longa et 2 mm lata. Corolla 12-13 mm longa, 17-18 mm diametro, alba; tubus 12—14 mm longa, 17-18 mm diametro, glaber, basi sub sepalis protrusa eaque occulta, apice in centro depresso; lobi 5-6 mm longi, 6—7 mm lati, apice leviter recurvi, intra breve papillati. Corona staminalis c. 8 mm longa et 10 mm diametro; apices loborum apicem styli1 aequantes. Columna staminalis c. 4 mm longa et 3 mm diametro; antherae appendices lanceolatae c. 1 mm longae et 0.7 mm latae; fissura alaris 0.8—1 mm longa sub antheris non prolongata. Apex styli c. 1 mm longus et 1 mm diametro. Pollinaria c. 0.96 mm longa et 0.77 mm lata; pollinia 0.77-0.78 longa, 0.29-0.31 mm lata; corpusculum 0.38-0.43 mm longum, 0.21—0.24 mm latum; caudiculae 0.11—0.14 mm longa, 0.5-0.9 mm lata. latiores in extremo corpusculari. Typus: Cultivated at Emerald Creek, Mareeba (from plant collected Papua New Guinea. MADANG Province: Headwaters Dom River, 4°58’S, 145° 45’E), 26 Nov 1995, IM Liddle IML1076 (holo: BRI |1 sheet + spirit]).
Wiry twiner; indumentum of uncoloured simple multicellular trichomes. Stems to several metres long, rooting at and between nodes, with sparse trichomes; internodes up to 200 mm long and 2 mm diameter, lenticillate with age. Leaf petioles cylindrical, 8-12 mm long, 1.5—3.5 mm diameter, faintly grooved along top, with sparse antrorse trichomes; colleters rounded, | or 2 at lamina base. Leaf lamina elliptic to elliptic- ovate, fleshy, up to 120 mm long and 60 mm wide; tip acute to shortly acuminate; base cordate with the lobes overlapping; venation of 6 or 7 looping lateral veins and indistinct reticulate minterlateral veins; upper surface green, venation + obscure, with scattered trichomes; lower surface pale green, venation indistinct,
Ht Pe CO ae SD ty ee ee ae ce De a ene oe ee a a* wane
Forster, Liddle & Liddle, Madangia inflata
with scattered to sparse trichomes. Flowering peduncles 8—22 mm long, 1.5—2 mm diameter, with scattered trichomes, with up to 9 flowers; bracts ovate-truncate, 0.4—0.5 mm long, 0.5—0.8 mm wide, sparsely ciliate. Flowers 12-14 mm long, 17—18 mm diameter; pedicels 32-45 mm long, 1.4—-1.5 mm diameter, glabrous. Sepals triangular, c. 1 mm long and 2 mim wide, glabrous. Corolla 12-13 mm long, 17-18 mm diameter, white; tube 12-14 mm long, 17-18 mm diameter, glabrous, base protuding down below sepals and obscuring them, top depressed in centre; lobes 5—6 mm long, 6—7 mm wide, slightly recurved at top, shortly papillate inside. Staminal corona c. 8 mm long and 10 mm diameter; lobe tips level with style-head. Staminal column c. 4mm long and 3 mm diameter; anther appendages lanceolate, c. | mm long and 0.7 mm wide; alar fissure 0.8—]1 mm long, not continuing down below anthers. Style-head c. 1 mm long and 1 mm diameter. Pollinaria c. 0.96 mm long and 0.77 mm wide; pollinia 0.77-0.78 mm long, 0.29-0.31 mm wide; corpusculum 0.38-0.43 mm long, 0.21—0.24 mm wide; caudicles 0.11—0.14 mm long, 0.5-0.9 mm wide, wider at corpusculum end. Fig. 1.
Additional speciinens examined: Known only from the type collection and live plants in cultivation.
Distribution and habitat: Madangia inflata is known only from the single type collection. Plants grow as twig epiphytes in the canopy of lowland complex mesophyll vineforest on volcanic soil. Hoya kenejiana Schltr. and miscellaneous ferns and orchids grow in association. Plants recorded in the surrounding forest include Tabernaemontana aurantiaca Gaud., 7. pandacaqui Lam., Aglaia sp., Graptophyllum pictum (..) Griff, Pittosporum sp., Popowia sp., Casearia sp., Rinorea horneri (Korth.) O.Kuntze, Maniltoa psilogyne Harms and Polyalthia sp.
Notes: Madangia inflata 1s unique in the Marsdenieae for its globose corolla and the form of the staminal corona. Globose corollas are rare in the Marsdenieae, the only regional example being the Australian Gunnessia pepo P...Forst. (Forster 1990). Madangia inflata does not appear closely allied to Gunnessia pepo
55
which has a totally dissimilar staminal corona and pollinaria. Globose corollas are more common, but still rare, in the Stapelieae (sensu Bruyns & Forster 1991), with convergently similar examples to Madangia being found in Echidnopsis malum (Lavranos) Bruyns (Bruyns 1988) and Stapeliopsis neronis Pillans (Bruyns 1981), both stem succulents from Africa. Unlike Echidnopsis malum, Gunnessia pepo and Sfapeliopsis neronis, the staminal column of Madangia inflata is not further enclosed by a tubular staminal corona within the enclosing corolla.
Madangia inflata is undoubtedly most closely allied to Hoya as it has a similar habit, and apart from the globose corolla and staminal corona, has a similar floral morphology to many species especially those in Papuasia. Globose corollas are extremely uncommon in Hoya with the only known example being H. heuschkeliana Kloppenburg from the Philippines. The corolla lobes in Hl. heuschkeliana are not as completely fused as in M. inflata with an obvious suture along the margins of each lobe, nor are they anywhere as large. Apart from its globose corolla, H. heuschkeliana has a staminal corona typical of Hoya.
The recently described H. telosmoides Omlor from Sabah is also worth mentioning here. This species has a corolla with a markedly inflated tube that has a constricted mouth and narrow lobes (Omlor 1996), These characters are not found in other species of Hoya, but H. telosmoides has a similar staminal corona to other species of that genus and Omlor (1996) concluded that it would be best placed there until such time as a monograph of Hoya could be undertaken and the status of the infrageneric classification reviewed.
The staminal corona of M. inflata differs markedly from that of Hoya. In Hoya the staminal corona comprises five separate lobes that are attached to the backs of the anthers fused into the staminal column. The lobes are discrete and although well separated in most taxa of Hoya may be closely contiguous in some species. The lower edge of each staminal corona lobe in Hoya is strongly inrolled along the sides
56 Austrobaileya 5(1): 53-57 (1997)
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oat a vasst * eewitu gem ere bee - s
5mm 5mm
Fig. 1. Madangia inflata. A. habit of flowering stem x 0.5. B. side view of flower x 3. C. face view of flower x 3. D, vertical cross-section of lower part of corolla tube, staminal corona and staminal column x 3.5. E, side view of staminal corona and staminal column x 3.5. F, face view of staminal corona and staminal column ~ 3.5. G. pollinartum x 25.
All from live and spirit material of D.J.Liddle IML1076. Del. B.Gray.
Forster, Liddle & Liddle, Madangia inflata
forming the ‘coronal groove’ of Rintz (1978). In Madangia the five staminal corolla lobes are contiguous to each other for almost their entire length, apart from the upper apex near the style head and the strongly recurved lower edges. The lower edge of each staminal corona lobe in Madangia is recurved at the end, although not at the sides, and forms a membranous frill, rather then a fleshy roll, that 1s positioned between involutions of the corolla wall. As a result no ‘coronal groove’ is formed. The only Hoya that approaches the staminal corona form of Madangia is H. multiflora Blume, that 1s sometimes segregated in the genus Centrostemma as C. multiflorum (Blume) Decne. In H. multiflora the staminal corona lobes do not have a marked ‘corona groove’, although it is still present, and the lower edge of the lobes are sharpely pointed (Rintz 1978). H. multiflora is a shrubby plant that never twines, not an uncommon trait in Hoya, and has a strongly reflexed rotate corolla. It is feasible that the floral features of a Hoya such as H. multiflora are plesiomorphic and similar to a putative ancestor to Madangia.
The pollinaria of Madangia inflata appear similar to those illustrated as group B of Malaysian Hoyas by Rintz (1978), where the caudicles are unwinged and the outer edge of the pollinia have a pellucid germination mouth.
The features of a globose corolla with totally fused corolla lobes, apart from the tips, and contiguous staminal coronal lobes that have a strongly recurved, frilled lower edge are considered derived for Madangia. Description of Madangia does not lend justification for dismemberment of Hoya into segregate groups as has been alluded to on occasion (e.g. Hill 1988). Despite the great morphological variability encompassed within the latter, the species presently recognised within Hoya are united by the shared character states of the staminal corona as described above.
Conservation status: The lowland rainforests of Madang Province are being cleared at an alarming rate for saw-logs and woodchips. The type locality for MZ inflata was a remnant of forest in an almost totally logged (clear-felled) area. Survival of this species in the area of its initial collection is unlikely 1n the near future.
ay
Etymology: The specific epithet is derived from the Latin inflatus (bladdery, thin, membranous and swollen) and alludes to the corolla texture and shape.
Acknowledgements
We are grateful to Bruce Gray of Atherton for the illustrations and Les Pedley (BRI) for the Latin translations. Fieldwork mm Papua New Guinea was possible due to a Christensen Research Institute Fellowship in 1992 and the then Director of CRI Dr M. Jebb assisted in various ways.
References
Bruyns, P.V. (1981). A review of Pectinaria Haw., Stapeliopsis Pillans and a new genus Ophionella (Asclepiadaceae). The Cactus and Succulent Journal of Great Britain 43: 61-83.
— (1988). A revision of the genus Echidnopsis Hook.f. (Asclepiadaceae). Bradleya 6: 1-48.
Bruyns, P.V. & Forster, P.I. (1991). Recircumscription of the Stapelieae (Asclepiadaceae). Taxon 40:381-391.,
Forster, P.I. (1990). Notes on Asclepiadaceae, 2. Austrobaileya 3: 273-289.
Hitt, K.D. (1988). A revision of Hoya (Asclepiadaceae) in Australia. Tefopea 3: 241-255.
Om or, R. (1996). Notes on Marsdeniecae (Asclepiadaceae) - a new, unusual species of Hoya from Northern Borneo. Novo 6: 288-294,
Rintz, R.E. (1978). The Peninsular Malaysian species of Hoya (Asciepiadaceae). Malayan Nature Journal 30: 467-522.
Marsdenia jensenti (Asclepiadaceae: Marsdenieae), a new Species from north-eastern Queensland
Paul I. Forster
Summary
Forster, Paul I. (1997). Adarsdenia jensenii (Asclepiadaceae: Marsdenieae), a new species from north-eastern Queensland. Austrobaileya 5(1): 59-61. The new species Adarsdenia jensenii is described and illustrated. It is endemic to north-eastern Queensland in the ‘Wet Tropics’ region and apparently restricted to a smal! area near Gadgarra State Forest.
Keywords: Asclepiadaceae, Marsdenieae, Marsdenia - Australia, Afarsdenia hemiptera,
Marsdenia jensenii.
Paul I. Forster, Queensland Herbarium, Meiers Road, Indooroopilly, Queensland 4068, Australia
Introduction
A revision of Marsdenia R.Br. in Australia and Papuasia was recently published wherein some seventy-eight species were enumerated (Forster 1995). The following new species was discovered too late to be included in that work, or in the subsequent Flora of Australia account (Forster 1996), and is described here.
‘Taxonomy
Marsdenia jensenti P.1.Forst., sp. nov. affinis M4. hemipterae H.Rch. latice albo (contra pellucido usque ad viridiflavo), florrum pedicellis 15—25 mm longis (contra 5—13 mm), corollae lobis valde dextris (contra + rectis usque = ad modice dextris), sepalis orbicularis usque ad orbicularo-ovatis (contra lanceolatis), coronae staminalis lobis apicem styli superantibus (contra non superantibus), poliniis ellipsoideis magnioribus 0.72—0.74 mm longis (contra globoso-ovoideus 0.5—0.55 mm longis) differt.
TFypus: Queensland. Coox District: State Forest Reserve 310, Gadgarra, track to Yeti Ridge, L7°17’°S, 145°44°E, 15 Feb = 1995, R. Jensen 105 (holo: QRS; iso: BRI). Canopy liane; latex white; indumentum of simple, multicellular trichomes that are uncoloured or
Accepted for publication 20 June 1997
pale golden in colour, Stems woody, lenticillate and up to 5 cm thick with age; internodes up to 150 mm long and 4 mm diameter, with scattered trichomes when young. Leaves petiolate; petioles 30-50 mm long, I-1.5 mm diameter, grooved on top, with scattered trichomes; colleters rounded to shortly subulate, 6 to 8 at base of lamina. Leaf lamina lanceolate-ovate to ovate, up to 160 mm long and 70 mm wide; tip acute to shortly acuminate; base weakly cordate to almost truncate; venation comprising 9 to 11 primary lateral veins per side of midrib and secondary interlateral reticulate veins; upper surface dark green, primary venation visible but not prominent, glabrous; lower surface pale green, primary and secondary venation prominent, glabrous. Inflorescence racemiform, up to 25 mm _ =4Iong; peduncle 11-15 mm long, c. 1 mm diameter, with sparse trichomes; bracts lanceolate to triangular, 1—1.5 mm long, 0.5—0.7 mm wide, with sparse to dense trichomes; flowers 1—20. Flowers 7-8 mm long, 17-22 mm diameter; pedicels 15— 25 mm long, |—2 mm diameter, with sparse trichomes. Sepals orbicular to orbicular-ovate, overlapping, 5.5—-6 mm long, 4—5 mm wide, with scattered trichomes, ciliate. Corolla rotate, 17— 22 mm diameter, green outside, faintly purplish inside; tube 1.5—2 mm long, not pouched at top, mainly glabrous but with a few scattered trichomes; lobes lanceolate with a bluntly acute tip, dextrorse, 7-10 mm long, 4.8—-S mm wide, mainly glabrous with a few scattered trichomes towards base, ciliate.
59-61 (1997)
Austrobaileya 5(1)
60
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Fig. 1. Adarsden
B—D from Jensen 105 (BRI). Del. W. Smith.
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SLC DA Ah cpa a a aa aR ee ee A eee
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Forster, Adarsdenia jensenii
Corolline corona absent. Staminal corona 4-5 mm long, 4-5.5 mm diameter, yellow; lobes in close proximity to one another, tips overtopping style-head, 7-10 mm long, 4.8—5 mm wide, upper portion lanceolate with an irregularly blunt tip, lower portion + rectangular. Staminal column c. 4mm long and 3 mm diameter; anther appendages ovate, 0.8~1 mm long, 0.8—1 mm wide; alar fissure c. 2 mm long. Style-head flattened-umbonate, c. 1 mm long, 2.5-2.8 mm diameter. Pollinaria c. 0.67 mm long and 1.53 mm wide; pollinia ellipsoid, 0.72—0.74 mm long, 0.34-0.36 mm wide; corpusculum oblong-ovate, 0.34—0.38 mm long, c. 0.19 mm wide; caudicles c. 0.95 mm long and 0.43-0.9 mm wide, widest at corpusculum end. Fruit cylindrical-oblong, c. 14.5 cm long and 5.5 cm diameter; seeds not seen. Fig. 1.
Additional spectnen examined : Queensland. Cook District: S.F.310 Gadgarra, track to Yeti Ridge, Jul 1995, Forster 17132 & Jensen (BRI).
Distribution and habitat: Marsdenia jensenti is thus far known only from Gadgarra State Forest at the western base of Bartle Frere, in the “Wet Tropics’ of north-eastern Queensland. Plants grow as canopy lianes in complex notophyill vineforest on soils derived from basalt.
Notes: Marsdenia jensenii appears to be related to M. hemiptera H.Rch. in general habit and floral morphology and keys to that species in Forster (1995). M. jensenti is easily differentiated from M. hemiptera on floral characters with its pedicels 15—25 mm long (versus 5—13 mm), strongly dextrorse corolla lobes (versus + straight to slightly dextrorse), orbicular to orbicular-ovate sepals (versus lanceolate), staminal corona lobes overtopping the style-head (versus not over-topping), and larger ellipsoid pollinia that are 0.72—0.74 mm long (versus globose-ovoid pollinia that are Q.5—0.55 mm long). The nearly glabrous foliage and white latex of M. jensenii also differs from that of M. hemiptera which is always beset with ferrugineous (ginger) indumentum and has clear or greenish-yellow latex.
61
Conservation status: Marsdenia jensenti appears to have a distribution largely restricted to Gadgarra State Forest where it appears to be reasonably abundant, as seedlings were common in places. It is highly likely that a detailed survey of the area will reveal further populations and despite the current paucity of herbarium records it 1s felt that no conservation coding is warranted.
Etymology: The specific epithet honours Rigel Jensen of Malanda who first discovered and collected this species.
Acknowledgements
Thanks to Rigel Jensen for showing me the plant in habitat, Les Pedley (BRI) for the Latin translation, and Will Smith (BRI) for the illustrations.
References
Forster, P.I. (1995), Circumscription of Afarsdenia (Asclepiadaceae: Marsdenieae), with a revision of the genus in Australia and Papuasia. Australian Systematic Botany 8: 703-933.
—— (1996). Marsdenia (Asclepiadaceae). In A.G.L. Wilson (ed.), Flora of Australia 28: 245-267. Melbourne: CSIRO Publications.
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A conspectus of Astrotricha DC. (Araliaceae) in Queensland, including two new species
A.R. Bean
Summary
Bean, A.R. (1997). A conspectus of Astrotricha DC. (Araliaceae) in Queensland, including two new species. Austrobaileya 5(1): 63-69. The eleven species of Astrotricha DC. indigenous to Queensland are listed in systematic order, with synonymy, place of publication and distribution given. Two new species, A. brachyandra and A. umbrosa, are described and illustrated. A comprehensive key to Queensland species is included.
Key words: Astrotricha, Araliaceae, key, taxonomy, Astrotricha brachyandra, Astrotricha umbrosa, Australian flora, Queensland
A.R.Bean, Queensland Herbarium, Meiers Road, Indooroopilly, Queensland 4068, Australia
Introduction
Astrotricha is an endemic Australian genus with representatives in Queensland, New South Wales and Victoria, and one species in Western Australia. Recent research has resulted in the description of several new species (Bean 1991, Makinson 1991, Bean 1995).
The present paper aims to summarise distributional and nomenclatural data on the species indigenous to Queensland, and to present descriptions of two new species.
No new infrageneric classification 1s proposed, but the published name A. subg. Hexocenia Domin 1s used for the species alhied to A. pterocarpa.
‘Taxonomy A key to Queensland species of Astrotricha DC. bre CAV OS] SATIS. as « sire bt rd sate oP oe dal Vise WANA Lech Ft 2 lee tees CEA EM AEA) og HEM cal eal lesb 2 LEA VCS. Soe lS PTV IGE ech ests bols ate eta) Genin: & SLATER ee hereew eH ETR lle dey doa adele eid cad 7 2. Petioles 5.5—21 cm long; inflorescence > 70 cm long; fruits winged ...............5. 3 Petioles 0.6—5.0 cm; inflorescence 20 —40 cm long; fruits without MATE Seg, axakinn BAU hase eiearsd aoe ED ALD bey a Meacin ion Atop ce ce SR Pace CELE A AG EG A ee ame 5 3, Leaves 9-16 cm wide, base cordate, midrib rounded below .............05. 3.A. cordata Leaves 1.7—5.5 cm wide, base obtuse, midrib keeled below. . 0... 0.0.00. cee eee 4 4. Inflorescence woolly-hairy, umbels 10—15-flowered, flowers creamy-white; leaves 1.7-4.0 cm wide... ... cee es 1.A. pterocarpa Inflorescence glabrous, umbels 7—1 1-flowered, flowers purple; leaves DOTA WIA! a Rd a, dhe woe pale Gasauleny WBunacl ates, ayise Chi Acerca arias 2.A. intermedia >. Flowers purple; stems mostly unbranched ....... 0... cee ee eee 4, A. roddii Flowers white or greenish-white; stems branched ........ 0.0. cece eee eee ees 6
__, Accepted for publication 9 July 1997
64
Austrobaileya 5(1): 63-69 (1997)
6. Petioles2.0-5.0cmlong ..... Ugrocat ras nd Sere eh anh dBaihidhas deen eeg el Sm tgPe dinar elk Soe 11. A. latifolia Petiones 0:04 3 CRON 0 doers dk eee dah tee ke ee ghana es 10. A. umbrosa Pee PISAVOS IGS Set vAlh ATTIC 2s revel ase Spcdestete os aeice: fs vee oaths oh Rd MOS Wie, acannaen eas ESE gee ais 8 LPaves G99 LS IWS oy, yo gi ane eeMig wate eo Ver on ae ee ROT a arb Aad Ge ol ele he ee 9 8. Leaves 3.5—7 mm wide, petioles 4-6 mm long; inflorescence 16 —30 cm PORN Serre oe yaces gooe, Sigens. Sag, ¥ deo ee oe Bhs ie 6a Se else EE OAS BUR dae Prep enyfelee CE yale Cea 6.A. biddulphiana Leaves 2.2—3.5 mm wide, petioles 3— 4 mm long; inflorescence 3—9 cm POTD. AB prsrcen hater ea ore etene cay me Aree ner eaere docevaced ane egane-emede: & 7.A. pauciflora 9, Leaves glabrous on both surfaces ...... 0... ccc cc eee eens 8.A. glabra Leaves with dense stellate tomentum on abaxial surface 2.0.0.2... 000. ee eee 10 10. Petioles (15-)23-30 mm long; flowers purple .............. 0000 ee eee 5.A. brachyandra Petioles 3—15 mm long; flowers white 2.0.0.0... ccc cece eee eee eee ee ens I] 11. Tomentum on stems and leaves loose, individual hairs c. 1 mm diameter; leaves 11~20 mm wide; pedicels 3-9.5 mmlong............ 000 eee eee 10. A. umbrosa Tomentum on stems and leaves tight, individual hairs c, 0.5 mm diameter; leaves (6-)7—11 mm wide; pedicels 8-15 mmlong ............ 9, A. longifolia
Astrotricha sabg. Hexocenia Domin, Biblioth. Bot. 89: 485 (1928). Type: Astrotricha pterocarpa Benth.
Monopodial shrubs, monocarpic; mericarps 3-locular, fruits winged (1n most species).
1. Astrotricha pterocarpa Benth., FI. Austral. 3: 379 (1867). Type: Queensland. Cook District: Fitzroy Island, WAills.n. (holo: Kx).
A description of this species is given in Bean (1991).
Distribution and habitat: A. pterocarpa is confined to north Queensland where it extends from the Melville Range area north of Cooktown southwards to Pentland and near Mackay. It grows in shallow sandy soil, over sandstone or eranite, in eucalypt woodland.
2. Astrotricha intermedia A.R.Bean, Austrobaileya 3(3): 524 (1991). Type: Queensland. LEicHHARDT District: Nogoa River camping area, Salvator Rosa National Park, 9 December 1989, R.O. Makinson 600 (holo: BRI; iso: CANB,
NSW).
Tlustration: Williams, Native Pl. Queensl. 3:31 (1987), as.A. pterocarpa.
Distribution and habitat: A. intermedia occurs along the central Queensland coast between Marlborough and Rosedale, and west almost as far as Tambo. It grows in open eucalyt woodland, in shallow sandy soils derived from sandstone, granite or sometimes serpentinite.
3. Astrotricha cordata A.R.Bean, Austrobaileya 3(3): 523 (1991). Type: Queensland. LEICHHARDT DISTRICT: Blackdown Tableland, c. 35 km SE of Blackwater, September 1971, R.J. Henderson H1018, £.Durrington & P. Sharpe (holo: BRI).
Illustrations: Williams, Native Pl. Queensl. 2: 47 (1984), as A. pterocarpa.
Distribution and habitat: A. cordata 1s sporadically distributed in southern Queensland from Blackdown Tableland to Mundubbera and Biggenden, and west to Salvator Rosa N.P. There is one known population in New South Wales, north-west of Grafton. It inhabits skeletal soils derived from sandstone, granite or occasionally rhyolite, in open eucaypt woodland.
2 |
Bean, Astrotricha
4. Astrotricha roddii Makinson, Telopea 4(2): 313 (1991). Type: New South Wales. NORTH WESTERN SLOPES: Macintyre Falls, 3 km S of junction of Macintyre and Severn Rivers, 23 November 1984, 4.N. Rodd 4096 (holo: NSW; iso: BRI, MEL).
Ilustrations: Flora of N.S.W. 3: 82 (1992).
Distribution and habitat: A. roddii 1s mainly distributed on the north-western slopes of New South Wales, particularly around Ashford. There is one population in Queensland, on Mt Bullaganang north-east of Texas. It occurs in shallow sandy acidic soils, with outcropping rocks, in woodland often dominated by Eucalyptus dealbata Cunn. ex Schauer and FE. melanophioia ¥.Muell.
Note: Although A. roddii does not possess the winged fruits characteristic of the other species of A. subg. Hexocenia, its placement there is supported by its unbranched slender habit and its 3-locular mericarps.
Astrotricha DC. subg. Astrotricha
Branched shrubs, perennial; mericarps 1-locular, fruits not winged.
5. Astrotricha brachyandra A.R.Bean sp. nov., affinis A. biddulphianae autem petiolis multus longioribus, foltis latioribus apice acuminatis, staminibus brevioribus, partibus omnibus inflorescentiae tomento stellato obsitis differt. Typus: Queensland. Port Curtis District: 41 km east of Biloela, in Kroombit Tops S.F., 24°25’S, 150°55’E, 15 November 1995, EF.J/. Thompson BIL207 & R.J.Price (holo: BRI; iso: CANB, MEL, NSW, distribuendt).
Shrub to 2 m high, erect, sparsely branched. Stems, petioles and abaxial surfaces of leaves stellate-pubescent, with individual hairs c. 0.1 mm across. Leaves alternate, narrowly lanceolate, 90-140 x 7-13 mm, acuminate, dark green adaxially, pale yellow abaxially, midrib impressed adaxially, raised abaxially; petioles terete, (15-)23—30 mm long. Panicles terminal, up to 60 cm long, densely
65
stellate-hairy, grey in colour; bracts ensiform, 1.0—2.5 mm long. Unit inflorescence umbellate, 5—9-flowered; peduncles and pedicels slender, pedicels 8—20 (-25) mm long. Flowers bisexual, protandrous, 4—5 mm across. Hypanthium stellate-hairy, 1.2—-1.9 mm long at anthesis, campanulate; calyx lobes deltate to semi-orbicular, apex acute or obtuse, c. 0.5 x 0.5 mm, purple, tomentose. Petals 5, deltate, 2.0-2.9 x 1.1-1.5 mm, purple, recurved, caducous; glabrous on adaxial surface, sparsely stellate hairy on abaxial surface; apex acuminate. Stamens 5, alternating with the petals; filaments terete, c. 0.6 mm long; anthers white, c. 1.4 mm long, versatile, dorsifixed, opening by longitudinal slits. Styles 2, 1.0-—2.0 mm long; at first erect, later spreading from base, persistent. Ovules | per carpel. Fruit a schizocarp, elliptical or somewhat quadrangular in transverse section, 5.0-6.5 mm long, 2.5-3.5 mm wide, 2-3 mm thick, surface stellate-hairy; brown to grey, splitting at maturity; mericarps 1-locular. Fig. 1 G—K. .
Spectinens exantined: Queensland, Port Curtis District: Kroombit Tops SF (boundary) on ridge above Annies gorge, Dec 1994, Brushe JB611 (BRI); creek flowing into Kroombit Ck near Annies Gorge falls, c. | km SE of falls, Oct 1995, Brushe JB201 (BRI); Kroombit Tops N.P., western section, Aug 1995, Thompson BILI14 & Turpin (BRI); 41 km E of Biloela, in Kroombit Tops S.F., Oct 1995, Thompson BIL181 & Turpin (BRI, CANB, MEL, NSW); 21.5 km NE of Biloela, Feb 1996, Thompson BIL214 & Price (BRI).
Distribution and habitat: A. brachyandra 1s known only from Kroombit Tops east of Biloela and the Callide Range north-east of Biloela. It grows in sandy soils derived from sandstone, as a component of woodland dominated by Eucalyptus trachyphloia F .Muell., £. suffulgens L.A.S.Johnson & K.D.Hill or &. cloeziana F.Muell. Altitudes vary between 600 and 800 metres.
Phenology: Flowers have been recorded from
October to December.
Affinities: A. brachyandra 1s closest to A. biddulphiana, and shares with that species the purple flowers and obconical fruits. A. brachyandra differs by the petioles being (15-)23-30 mm long (4—6 mm long for
66
A. biddulphiana), leaves 7-13 mm wide with acuminate apex (3.5~7 mm wide with acute apex for A. biddulphiana), shorter stamens and presence of stellate tomentum on all parts of the inflorescence.
Conservation status: The risk category for Astrotricha brachyandra according to the criteria of Chalson & Keith (1995) is “priority for investigation’ (criterion a), or 2RC1 according to the system of Leigh and Briggs (1996). The species is known only from Kroombit Tops and Callide Range, with a total geographical range of about 50 kilometres. The recommended conservation status for this species as defined by the Queensland Nature Conservation Act 1992 ts ‘rare’.
Etymology: The species epithet brachyandra is derived from the Greek brachy-short and andros-male, and refers to the short stamens which are a feature of this species.
6. Astrotricha biddulphiana F.Muell., Victorian Naturalist 7: 115 (1890). Type: Queensland. MARANOA DISTRICT: near Mt Playfair, H. Biddulph s.n. (holo: MEL? n.v.).
Astrotricha ledifolia var. glabriflora F.M.Bailey, Queensland Flora 2:732 (1900). Type: Queensland. near Rockhampton, undated, O’Shanesy s.n. (holo: MEL, photo!).
Distribution and habitat: A. biddulphiana occurs in southern Queensland away from the coast, extending from Lake Elphinstone to Barakula S.F. near Chinchilla, and west to Bull Creek gorge and Mt Playfair. It grows in eucalypt woodland on shallow sandy soils over sandstone or granite.
7. AStrotricha pauciflora A.R.Bean, Austrobaileya 4(3): 407 (1995). Type: Queensland. Moreton Districr: North Ridge, Mount Barney N.P., 16 July 1994, A.R.Bean 7734 & D.A.Halford (holo: BRI; iso: CANB, K, MEL, NSW).
Distribution and habitat: A. pauciflora 1s confined to Mt Barney and Mt Maroon, both in the Mt Barney National Park, Queensland,
Austrobaileya 5(1): 63-69 (1997)
close to the N.S.W. border. It grows in rocky crevices in low eucalypt woodland or heathland, at altitudes above 600 metres.
8. Astrotricha glabra Domin, Biblioth. Bot. 89: 486 (1928); Astrotricha longifoliavar. glabrescens F.M.Bailey, Queensland Agricultural Journal 21(6): 293 (1908). Type: Moreton Island, 12—14 September 1908, C.T. White s.n. (ecto (here chosen): BRI [AQ 333042]).
Distribution and habitat: A. glabrais endemic to south-eastern Queensland and is found on North Stradbroke Island, Moreton Island, Fraser Island and the Cooloola area. It grows under eucalypt forest in deep sands of quaternary age.
Note: In his treatment of Astrotricha glabra, Domin (1928) attributes the original publication of a varietal epithet ‘g/abra’ to F. Mueller in Fragmenta Phytographiae Australiae 7: 148. While Mueller did discuss the occurrence of a glabrous form there, he did not provide a description of the plant, nor did he coin the epithet ‘glabra’. The correct authorship citation is hence A. glabra Domin.
9, Astrotricha longifolia Benth., in S.L. Endlicher et al., Enum. pl. 54 (1837). Type: New South Wales. Sydney area (syn: Bauer s.n.; Sieber 538).
Illustrations: Williams, Native PI. Queensl. 2: 47 (1984).
Distribution and habitat: A. longifolia inhabits coastal areas of southern Queensland and New South Wales. It grows in low woodland dominated by Eucalyptus spp. or Banksia spp., in sandy soils.
10. Astrotricha umbrosa A.R.Bean sp. nov., affinis A. fongifoliae autem tomento laxiore ex pilis stellatis majoribus constante, foliis plerumque 11-20 mm latis (non 6-11 mm latis), pedicellis 3—9.5 mm longis (non 8-15 mm longis) differt. Typus: Queensland. BURNETT District: Dead End lookout, Coominglah S.F., west of Monto, 17 August 1995, A.R.Bean 8859 (holo: BRI; iso: CANB, DNA, MEL, NSW).
pes in SSSR SIS Do Fd Rr PO I rs SERRE Teme ae eET
THEN See.
Jesexo Shhies.
LASSE ESE a
3 cee CaS a PR
Re ee Cate tt Ld reer el Ba braced
Bean, Astrotricha 7 67
Fig. 1. Astrotricha umbrosa A. portion of inflorescence x 4. B. flower at male stage x &. C. flower at female stage x 8. D. fruit x 4, E. leaf x 0.5. F. stellate hairs x 8. Astrotricha brachyandra. G. portion of inflorescence < 4. H. flower at male
stage x 8. I. flower at female stage x 8. J. fruit x 4. K. leaf x 0.5. A-C,E,F Bean 9110; D, Bean 2540; G-K, Thompson BIL207 & Price. |
68
Astrotricha sp. (Pomona A.R.Bean 1157) in Bean (1994).
Shrub to 2 m high, erect, well-branched. Stems, petioles and abaxial surfaces of leaves stellate-pubescent, with individual hairs c. 1.0 mm across. Leaves alternate, lanceolate to narrowly lanceolate, (50)70—120 x (8-) 11-20 mm, acuminate, dark green adaxially, pale yellow abaxially, midrib impressed adaxially, raised abaxially; petioles terete, 6-13(-17) mm long. Panicles terminal, 10-30 cm long, densely stellate-hairy, brown to yellow in colour; bracts ensiform, 1.7—3.5 mm long. Unit inflorescence umbellate, (4-)5—11-flowered; peduncles and pedicels slender, pedicels 3.0—9.5 mm long. Flowers bisexual, protandrous, 4.5—6.0 mm across. Hypanthium stellate-hairy, 1.0—1.5 mm long at anthesis, campanulate; calyx lobes deltate, apex acute or obtuse, c. 0.4 x 0.4mm, green to white, stellate-hairy or mostly glabrous. Petals 5, deltate, 1.52.5 x 0.7—1.1 mm, white or pinkish, somewhat persistent; glabrous on adaxial surface, stellate hairy on abaxial surface; apex acute to acuminate. Stamens 5, alternating with the petals; filaments terete, c. 1.9 mm long; anthers white, c. 1.1 mm long, versatile, dorsifixed, opening by longitudinal slits. Styles 2, 1.5-2.8 mm long; at first erect, later spreading from base, persistent. Ovules I per carpel. Fruit a schizocarp, + orbicular in outline, 3.3-4.0 mm long, 3.9-4.3 mm wide, 1.2—-1.5 mm thick, glabrescent; brown, splitting at maturity; mericarps |-locular. Fig. 1 A-F.
Specimens examined: Queensland. Port Curtis District: Kroombit Tops SF 316, 48 km E of Biloeia, Sep 1988, Gibson TOT431 (BRI); near headwaters of Cedar Ck, SF 583, Jul 1995, Thompson CAL312 & Turpin (AD, BRI, DNA, NSW). Burnett District: near Dead End Lookout, Coominglah SF, near Monto, Aug 1990, Bean 2086 (BRI, MEL, NSW); Cania Gorge NP, c. 600 m W of Mt Dowgo, Nov 1995, Brushe JB315 & Hendry (BRI); c. 15 km (direct) WNW of Monto, turn-off from Forest Drive to Dead End Lookout, Coominglah SF, Oct 1988, Adakinson 550 & Krauss (AD, BRI, CANB, CHR, K, MEL, NSW). Wipe Bay District: Mt Cooroora, near Pomona, Oct 1989, Bean 1157 (BRD; Linda Garrett Park, 2 km W of Mapleton, Oct 1990, Bean 2539 (BRI, NSW); Oakey Creek road, Mapleton SF, Oct 1990, Bean 2540 (BRI); Scenic Drive, Blackall Range, c. 5 km N of Mapleton, Sep 1988, Sharpe 4824 et al. (BRI, NSW). Moreron District: Delicia road, Mapleton, Dec 1989, Bean 1257 (BRI); Cooloolabin SF, Mapleton Forest Drive, 2 km N of Mapleton, Sep 1989, Leiper s.n. (BRD).
Austrobaileya 5(1): 63-69 (1997)
Distribution and habitat: A. umbrosa is endemic to southern Queensland, extending from Kroombit Tops and the Boyne Valley, to the Mapleton area. It grows in relatively dense and shady open forest dominated by Eucalyptus spp. including &. pilularis Sm., E. cloeziana F.Muell. and £. acmenoides Schauer, in sandy or sandy loam soils.
Phenology: Flowering occurs in September- October.
Notes: A. umbrosa 1s similar in appearance to A. floccosa, but it differs by its ensiform bracts 1.73.5 mm long (filamentous, 2-10 mm long for A. floccosa), inflorescences 10-30 cm long (30-100 cm long for A. floccosa) and the 5—l1-flowered umbels (12—20-flowered for A, floccosa).
From A. longifolia, it differs by its stellate hairs c. 1.0 mm across (c. 0.5 mm for A. longifolia), looser tomentum, leaves mostly 11-20 mm wide (6-l[lmm wide for A, longifolia), and pedicels 3.0—9.5 mm long (8-15 mm for A. /ongifolia).
Conservation status: The risk category for Astrotricha umbrosa according to the criteria of Chalson & Keith (1995) is ‘susceptible’ (criteria e and f), or 2RCi according to the system of Leigh and Briggs (1996). The species is known from about half a dozen widely scattered populations. The recommended conservation status for this species as defined by the Queensland Nature Conservation Act 1992 is ‘rare’.
Etymology: The species epithet is from the Latin wmbrosus meaning shady or growing in shade and refers to the habitat of the species in rather dense, shady forests.
11. Astrotricha latifolia Benth., in S.L. Endlicher et al., Enum. pl. 54 (1837); A. floccosa var. incana Benth., FI. Austral. 3: 380 (1867); A. floccosa vat. latifolia (Benth.) Domin, Biblioth. Bot. 89: 486 (1928), nom. illeg. Type: ‘in collectione Baueriana’ (? W n.v.).
Illustration: Flora of N.S.W. 3: 82 (1992),
VSM rrr cam an le DOE had ee ada nad te dee e tn ht ATED TEN eA DE
Bean, Astrotricha
Distribution and habitat: A. latifolia occurs in southern Queensland south from about Gympie and west to the Great Dividing Range, and along much of the New South Wales coast. It inhabits tall eucalypt forest in high rainfall areas.
Acknowledgements
I am grateful to John Thompson for his persistent efforts to collect good flowering and fruiting material of A. brachyandra, to Will Smith for the illustrations, and Les Pedley for the Latin diagnoses.
References
Bran, A.R. (1991). Notes on Astrotricha DC. (Araliaceae) in Queensland. Austrobaileya 3(3): 523-528.
(1994). Astrotricha, In R.J.F. Henderson (ed,) Queensland Vascular Plants: Names and Distribution. Brisbane: Queensland Department of Environment and Herttage.
(1995), A new species and new record for Astrotricha DC, (Araliaceae) in Queensland. Austrobaileya 4(3): 407-409.
CuHaLson, J.M. & Keitu, D.A. (1995). A Risk Assessment scheme for Vascular Plants: Pilot Application to the Flora of New South Wales, Hurstville: National Parks and Wildlife Service.
Domm, K. (1928). Astrotricha, In Bettrage zur Flora und Pflanzengeographie Australiens. Bibliotheca Botanica 89: 485-487,
Briaces, J.D. & Leicu, J.H. (1996). Rare or Threatened Australian Plants. Melbourne: CSIRO.
Makinson, R.O, (1991). Two new species of Astrotricha (Araliaceae) from New South Wales. 7e/opea 4: 313-319.
69
Cooktownta robertsii, a remarkable new genus and species of Orchidaceae from Australia
David L. Jones
Summary
Jones D.L. (1997). Cooktownia robertsii, a remarkable new genus and species of Orchidaceae from Australia, Cooktownia robertsii D.L. Jones is described from Queensland. Austrobaileya
5(1): 71-78.
Keywords: Orchidaceae - Queensland, Orchidoideae, Orchideae, Cooktownia robertsii
D.L. Jones, Centre for Plant Biodiversity Research, Environment Australia, Box 1600, Canberra,
Australian Capital Territory 2601, Australia
Introduction
Identification of the genus of an unfamiliar species of orchid becomes relatively straight forward with some experience. The generic placement of an extremely unusual terrestrial orchid recently collected in northeastern Queensland, however, has caused considerable difficulty. Embryological studies (Clements pers. comm.) show that the species has similar developmental stages to orchids of the subfamily Orchidoideae, tribe Orchideae (Abe 1972; Fredrikson 1991; Clements 1995). The elongated, somewhat fusiform, asymmetric ovaries, the shape and arrangement of the sepals and petals and the shape of the elements of the column are consistent with this placement. However the paired, broadly ovate, eround-hugging leaves and the thin wiry scape are very different from any Orchidoid species which has been described from the region. The labellum, which is entire, lacks a basal spur, usually considered to be a feature of orchids of this type, but Dressler (1993) notes that a spur may be lacking in some species.
Examination of the flowers of this novelty shows that the column is much modified and would appear to be non-functional. Elements of the anther and stigma occur as simple rod-like analogues (Fig 1.1). The anther analogue can be recognised by its dorsal placement opposite the labellum
Accepted for publication 15 August 1997
(Fig. 1.3 & 1.4) and the presence on its lateral surfaces of aggregates of structures which resemble pollen tetrads (Fig. 1.5, 1.6). Two or four rod-like structures anterior to the anther analogue are apparently much modified stigmas (Fig. 1.2) and/or rostellar arms (for developmental studies in Orchidoid species see Kurzweil 1987). Thus although the flowers of this enigmatic species appear to be supertficially normal, the column is much modified and tts elements would appear at first sight to be non-functional, lacking pollen and receptive stigmata. Despite these significant reproductive limitations the species 1s common over a localised natural area and is maintaining a viable population level (L.Roberts pers. comm., C.Broers pers. comm.). Examination of mature ovaries shows that they contain reproductive structures which have an appearance remarkably similar to normal Orchidoid seed (Fig. 2).
Subtribal Placement
The subtribal classification of the Orchideae is difficult and well beyond the scope of this paper. A brief consideration of the subject 1s necessary since the subtribal characters influence the generic choices for the taxon in question. Dressler (1993) recognised two subtribes within the Orchideae, the Habenariinae and Orchidinae, and at the same time noted that the distinction between them needs reevaluation. In Orchidinae, according to Richard (1818) and Dressler (1993), the
ustrobaileya 5(1): 7i~—78 (1997)
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