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| O R I G I N A L A R T I C L E Life-history characters and phylogeny are correlated with extinction risk in the Australian angiosperms A. Sjo¨stro¨m 1 and C. L. Gross 2 * 1 School of Mathematics, Statistics and Computing Science and 2 Ecosystem Management, School of Environmental Sciences and Natural Resources Management, University of New England, Armidale, NSW, Australia *Correspondence: C.L. Gross, Ecosystem Management, School of Environmental Sciences and Natural Resources Management, University of New England, Armidale, NSW, 2351, Australia. E-mail: cgross@une.edu.au These authors contributed equally to this work. A B S T R A C T Aim To determine whether life-history characters that affect population persistence (e.g. habit and life span) and those that influence reproductive success (e.g. sexual system and fruit type) are non-randomly correlated with extinction risk (i.e. threat category) in the Australian flora (c. 19,000 species, of which c. 14% is threatened). To identify patterns that present useful conservation directions. To understand patterns of extinction risk in the Australian flora at a broad scale. Location Continental Australia. Methods A country-wide exploration of four life-history characters in the Australian flora (n ¼ 18,822 species) was undertaken using reference texts, expert opinion, herbarium records and field work. For each character and threat- category combination, a G-test (using a log-linear model) was performed to test the null hypothesis that the two factors were independent in their effects on count. A generalized linear model (GLM) with a logit link and binomial error distribution was constructed with the proportion of taxa in each extinction risk category as the response variable and the habit, sex and fruit-type characters as explanatory terms. In a separate approach, we investigated patterns across the threat categories of non-endangered extant, endangered, and extinct using a multinomial model. We examined whether or not species-poor genera were more likely to contain threatened or extinct species than species-rich genera. A GLM with a binomial error distribution and logit link function was constructed to obtain a weighted regression on the proportion of species listed as extinct or endangered within a genus versus the log of the size of the genus. We also used a supertree analysis and character tracing to investigate the role of phylogeny on extinction risk. Results We found that the Australian flora is primarily composed of bisexual shrubs with dry-dehiscent fruits. Dioecious breeding systems (separate female and male flowers on separate plants) in many floras are the predominant unisexual system, but in Australia there are unexpectedly high levels of monoecy (separate female and male flowers on the same plant). Within the extinct data set of 31 species we detected a significant departure from that expected for habit but not for life span, sexual system or fruit type. There are significantly fewer trees on the extinct list than expected. This may reflect the greater resilience of trees than of other growth habits to extinction processes as well as the observation time-frame. Within the endangered data set of 450 species we found significant differences in the representation of the observed characters from that expected within sex systems and fruit types. We show that, depending on the life form, unisexual breeding systems can be significantly and positively associated with endangered species compared with non-threatened species. For example, there are more Journal of Biogeography (J. Biogeogr.) (2006) 33, 271–290 ª 2006 The Authors www.blackwellpublishing.com/jbi 271
Journal compilation ª 2006 Blackwell Publishing Ltd doi:10.1111/j.1365-2699.2005.01393.x
I N T R O D U C T I O N There are many different types of events that can increase extinction risks in species; for example, fragmentation has multiple effects on ecosystems (Laurance et al., 2002) and is a key factor worldwide that can prematurely halt population persistence. However, it is not always that a species was just in the wrong place at the wrong time – even related species in the same landscape can differ in their resilience to perturbations (e.g. Bertya ingramii versus B. rosmarinifolia, Scott & Gross, 2004). Are there inherent properties in species that predispose them to vulnerability or resilience? A broad approach to address this question is to determine whether or not there are specific traits clustered with the state of extinction risk. Increasingly this approach is being used to evaluate the properties of rarity and speciosity geographically or within lineages (e.g. Hegde & Ellstrand, 1999; Rey Benayas et al., 1999; Edwards & Westoby, 2000; Murray et al., 2002a,b; Golding & Hurter, 2003) in an attempt to forecast the types of species that may be vulnerable to extinction. The evolutionary significance of trait clustering can also be examined using correlation tests of life-history characters against ecological parameters (e.g. Chazdon et al., 2003). However, seldom is there a comprehensive data set that covers whole continents (see later). Consequently an overall appraisal of extinction risk is mostly lacking at continental scales. In most floras, for example, so little is known about the distribution of life- history characters on a continental scale that partial data sets (e.g. a clade, Murray et al., 2002a) are used instead, with the added power of phylogenetic analyses in some cases (e.g. Murray et al., 2002b; Murray & Lepschi, 2004). Biogeographi- cal attributes such as habitat type may also be important predictors of increased extinction risk; however, our initial focus is at the Australia-wide scale and on whether or not there are life-history attributes correlated with increased extinction risks. Perhaps the predominance of a life-history character (e.g. habit) is skewed in abundance in both threatened species and the greater population – it is difficult to resolve when data on the greater population are unavailable. The Australian vascular flora presents a challenge in this area: there are at least 19,000 vascular species yet the abundance and distribution of fundamental life-history characters, such as those that affect population persistence (e.g. habit and life span) and those that influence reproductive success (e.g. sexual system and fruit type), are poorly known on a continental basis. Such data, if available, could be used to examine the general occurrence of patterns of rarity in the flora. There is certainly a need for a broad approach as the estimate for the number of threatened vascular plants in Australia is alarmingly high at 14.4% (Walter & Gillett, 1998). Establishing the abundance and distribution of life-history characters for an entire flora is a challenge, yet knowledge of these properties at such a broad and inclusive scale could be a powerful tool for conservation planning. As a starting point we undertook a study of the distribution of four life-history characters that we consider fundamental to all species. These characters were habit, life span, sex system and fruit type, and they were examined for Australian recently extinct and endangered angiosperms and for the Australian flora as a whole. Habit (i.e. life form) has been linked to increased extinction risk in several floras (Robinson et al., 1994; Turner et al., 1996; Hegde & Ellstrand, 1999; Rogers & Walker, 2002) and it is intricately linked with life span (Garcia, 2003). Sex system can also be associated with elevated extinction risks, particularly for unisexual species. For example, Vamosi & Vamosi (2005) found that dioecious species experience higher extinction rates and (or) lower speciation rates compared with non-dioecious sister groups. They also found that the woody growth habit is probably a contributing factor to the higher incidence of dioecious species being at risk of extinction but that the character was not solely responsible for the pattern. Certain fruit types can be non-randomly associated with rare species (Hegde & Ellstrand, 1999) too, and as fruit type can be correlated with sex system and habit (reviewed in Gross, 2005) fundamental life-history characters such as habit, life span, sex system and fruit type should not be examined in isolation from each other. We use the information on life-history characters to describe first the composition of the Australian flora and then patterns in trait distribution. Next we determine the utility of taxon richness as a predictor of extinction (see Schwartz & monoecious species than expected by chance among the tree species listed as endangered but fewer among the herbaceous life forms. Threat category was found to be non-randomly clustered in some clades. Main conclusions Life-history characters in certain combinations are predictive of extinction risk. Phylogeny is also an important component of extinction risk. We suggest that specific life-history characters could be used for conservation planning and as an early warning sign for detecting vulnerability in lists of species. Keywords Australia, correlated evolution, extinction risk, flora, fruit types, genus size, life- history characters, phylogenetic analyses, sex systems. A. Sjo¨stro¨m and C. L. Gross 272 Journal of Biogeography 33, 271–290 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd Simberloff, 2001). Finally we compare our results for the Australian continent with other regions and provide informa- tion that could be incorporated into conservation planning. M A T E R I A L A N D M E T H O D S The number of extant, endangered and extinct flora in Australia Extant flora To begin we compiled a list of the flowering plant families for Australia (n ¼ 222 families) and determined the number of species in each family by consulting Hnatiuk (1990), and, when it became available, the online version, the Australian Plant Name Index: http://www.anbg.gov.au/cgi-bin/apni. We checked the number of species and revised the counts when more recent treatments became available, for example from the Flora of Australia volumes, regional floras, or other taxonomic treatments. Family names were ultimately delineated following the Angiosperm Phylogeny Group classification (Stevens, 2001 onwards). Endangered and extinct flora There is much discussion about the membership and robustness of threatened species lists (Akcakaya et al., 2000; Burgman, 2002; Possingham et al., 2002; Keith & Burgman, 2004). Our list of extinct and endangered species was derived from the schedules within the Environmental Protection and Biodiversity Act 1999 (EPBC Act) as of November 2003.* This group of species was considered by us to represent species most likely to be extinct or to go extinct in the next 10–20 years. We refer to these species as rare because they variously occur within the seven forms of rarity as proposed by Rabinowitz (1981). There are many thousands more species considered as rare in threatened species classifications in Australia (e.g. Briggs & Leigh, 1996), but these are generally not considered to be at a high risk of extinction in the next 10 years and are not considered separately in this study.
To verify the accuracy of Australian taxa listed as presumed extinct, we sought collection and taxonomic information for 61 taxa. In particular, an attempt was made to find the protologue, additional descriptive material, and the most recent sighting or collection date for each taxon presumed extinct. Only flowering species were considered. We then only listed a species as extinct if (1) it was taxonomically distinct (if the species is known only from a single specimen, it was placed in the ‘known only from the type’ category and not considered further), (2) no collections or sightings of the species are known within the last 50 years, and (3) the species is not extant outside Australia. No additional checking was performed on the validity of membership on the endangered list except to exclude non- vascular taxa and taxa below the species level or to revise a name according to the latest systematic information. Life-history characters In order to develop a set of testable hypotheses for investi- gating the significance of the trends observed within the extinct and endangered flora, against the extant flora as a whole, we assayed four life-history characters for 31 extinct species, 450 endangered species and for the Australian angiosperms, which totalled 18,821 species (including 1997 introduced species). We only included taxa at the species level. We looked at four life-history characters for each family: habit, life span, sex system and fruit type. Habit (i.e. life form) was categorized as herb, vine, liane, shrub, or tree. Life span was categorized as either short-lived (i.e. annuals or biennials) or long-lived (perennials). Sex system was categorized as hermaphroditic (i.e. all flowers bisexual on a plant), dioecious (i.e. female and male flowers found on separate individuals and includes andro- and gynodioecious species), or monoecious (i.e. female and male flowers found as separate flowers on the same individual and includes andro- and gynomonoecious and polygamomonoecious species). Fruit type was categorized as dry-indehiscent (including achene, caryopsis, nut, nutlet, samara, some schizocarps, utricle, and some capsules), dry- dehiscent (including most capsules, stroboli, follicles, legume, mericarp, saliqua, most schizocarps, and loments), or fleshy (including berry, drupe, gyconium, pome, pseudocarp, spathe, synangium, syncarp, viviparous seedlings, pipo, and pyrene). When all species within a family shared the same combi- nation of life-history characters (see below), the data were tallied at the family level for the number of species known to occur in the family. For example, if all species within a family were hermaphroditic, annual, herbs with capsules, the char- acter data were tallied along with a count for the number of species in the family and individual species were not looked at in detail. For families where categories varied within a life- history character (e.g. some species were hermaphroditic and other were dioecious), we looked at the life-history characters at the generic level and if these were consistent we tallied them; otherwise, we examined life-history characters for each indi- vidual species. To determine which category within a character a species exhibited, we sourced the information in the first instance either from the Flora of Australia volumes, or from Morley & Toelken (1983), or from regional botanical reference texts (a list is available from the authors). If we were not able to obtain the information from these sources, we gathered and queried information from published taxonomic treatments, researchers, protologues or herbarium specimens. For some species we were unable to determine every life-history char- acter. In these cases we omitted from the analyses that life- history character for that species (habit data were lacking for 129 spp., life span for 135 spp., sex system for 192 spp., and fruit type for 34 spp.). *The current EPBC Act list is available online at http://www.deh. gov.au/biodiversity/threatened/species/index.html. Life-history characters in the Australian flora Journal of Biogeography 33, 271–290 273 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd Statistical analyses Life-history characters Prior to comparative analysis of the non-endangered extant and endangered species, introduced species were excluded. Of the remaining species, each was coded individually according to its life-history characters and conservation status (i.e. non- endangered extant or endangered). Owing to the autocorrela- tion of habit and life span, only the characters habit, sex, and fruit type were considered. Analysis then proceeded with the follow- ing steps. The R statistical package (Ihaka & Gentleman, 1996) was used to obtain contingency tables of each separate character and conservation status. For example, a table was constructed of counts of species classified by habit (5 categories) and conservation status (endangered or non-endangered extant). For each character and status combination, a G-test (that used a log-linear model) was performed to test the null hypothesis that the two factors (i.e. individual characters and status) were independent in their effects on count (Crawley, 2003). Next, characters were considered together in a four-way frequency table (i.e. all character combinations and status). To model the data, conservation status (endangered or non- endangered extant) was reformulated as a proportion for ease of analysis (Crawley, 2003). A GLM (generalized linear model) with a logit link and binomial error distribution was constructed with the proportion of taxa as endangered as the response variable and the habit, sex and fruit-type characters as explan- atory terms. The saturated model consisted of the main effects, all two-way interactions, and the three-way interaction of explanatory terms. The saturated model was reduced by stepwise removal of terms that reduced the Akaike Information Criterion (AIC; Akaike, 1974). A similar procedure was followed to compare extant species and extinct species. As above, introduced species were excluded from the analysis. R was used to obtain a four-way frequency table. The response y (extinct or extant) was reformulated as a proportion for ease of analysis. A saturated GLM model with a logit link and binomial error distribution was constructed with the proportion of taxa extinct as the response variable, and the main effects, all two-way interac- tions, and the three-way interactions as explanatory terms. This model was reduced using the step function in R. In a separate approach we investigated patterns across status levels of non-endangered extant, endangered, and extinct species using a multinomial model (Venables & Ripley, 2002) that was fitted to the data with the response y as the conservation status level (extinct, endangered, or non- endangered extant) and explanatory terms as the life-history characters. As for the GLM models, the step function in R was used to reduce the saturated model. Species richness We examined whether or not species-poor genera were more likely to contain threatened or extinct species than species-rich genera. We excluded genera that only contained introduced species (n ¼ 660). For each genus (n ¼ 1978), we determined the proportion that was endangered or extinct relative to the overall size of the genus in Australia. R was used to investigate the influence of generic species richness on membership in the extinct species list. A GLM with a binomial error distribution and logit link function was constructed to obtain a weighted regression on the proportion of extinct species within a genus versus the log of the size of the genus. Similarly, a weighted regression was also performed on the proportion of endan- gered species versus the log of the size of the genus. In both cases, the significance of the parameter values was assessed by a deletion test in which the original model was compared with a simpler model that was fitted with a constant (i.e. the null model).
For both the extinct and endangered species data we explored the relationship with rarity further by using a second model that was constructed to include only those genera that contained extinct species (n ¼ 26) or endangered species (n ¼ 179), respectively. The reliability of the parameters estimated for the latter was assessed using a bootstrap test. The procedure followed was to construct a model that used the original endangered proportion data as the response variable. The explanatory term was the log of the size of the genus, and the number of genera considered consisted of a random sample (n ¼ 179) of all genera. A weighted regression was performed on each of 10 000 different random samples and the resulting parameter values compared. We repeated the procedure above to investigate familial speciosity and the presence of endangered or extinct species. In particular, we excluded families that contained only intro- duced species (n ¼ 20). For each family (n ¼ 202), we determined the proportion of taxa that was endangered or extinct relative to the overall size of the family in Australia. A GLM with a binomial error distribution and logit link function was constructed to obtain a weighted regression on the proportion of extinct species within a family versus the log of the size of the family. Similarly, a weighted regression was also performed on the proportion of endangered species versus the log of the size of the family. In both cases, the significance of the parameter values was assessed by a deletion test in which the original model was compared with a simpler model that was fitted with a constant (i.e. the null model). For both the extinct and endangered species data, a second model was constructed by including only those families that contained extinct species (n ¼ 16) or endangered species (n ¼ 62), respectively. The reliability of the parameters estimated for the latter was assessed using a bootstrap test as described previously. The association of rarity with phylogeny A supertree of the Australian flowering species was too large to work with and so three smaller trees (Asterids, Core Eudicots and Commelinids) were constructed using the Phylomatic database (Webb & Donoghue, 2002), an automated process A. Sjo¨stro¨m and C. L. Gross 274 Journal of Biogeography 33, 271–290 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd that builds a hypothetical tree based on a master phylogeny – we chose the angiosperm supertree from Davies et al. (2004), which is based on 46 source trees, as our master phylogeny. Genera were used as terminal taxa in this analysis because of the large number of poorly understood infra-generic lineages and as an attempt to minimize non-independence among related groups. Insufficient information on relationships between or within taxa is presented as polytomies. Branch lengths were not known for all relationships and were given unit length. A nexus version of these trees is available on request. The trait, not threatened or threatened (endangered or extinct), was traced as a binary character over each tree using macclade
version 4.03 (Maddison & Maddison, 2001). Thus genera containing an endangered or extinct species were given a score of 1 while genera without an endangered or extinct species were given a score of 0. Clustering of the trait (phylogenetic conservatism) was examined by determining the number of steps required for each character state (summed cost of all changes, gains and losses) in each of the three trees. Polytomies were set as soft to represent uncertain resolution and so that the polytomy is resolved in the most parsimonious way for that character. (Hard polytomies, on the other hand, assume, for any character exhibited among a group of taxa, that character has evolved independently for each taxon, i.e. simultaneous speciation events.) The character state was reshuffled among the taxa in 1000 random resolutions of each tree. For each tree, character tracing was then used to compare the actual number of steps with the number of steps in the 1000 trees based on randomly reshuffling the character states. If the actual number of steps ranked within the lowest 5% of the 1000 random resolutions, the character was considered significantly phylogenetically clustered. The concentrated-changes test in MacClade version 4.03 (Maddison & Maddison, 2001) was used to examine whether observed associations among traits (e.g. rarity and genus size) reflect phylogenetic correlations. This test examines whether gains (0 to 1 changes in a binary character) in one character (e.g. extinct status) are more concentrated than expected by chance on those branches of the tree that are reconstructed to have a particular distribution of state in the second character (e.g. genus size). This test can only be used on resolved trees (i.e. without polytomies), and thus 10 randomly resolved trees were used so that a range of probabilities could be reported. Threat status was traced against genus size [1 ¼ small (1–2 species) 0 ¼ or not small] and in a separate set of analyses against sex systems (unisexual or bisexual) and against fruit type. R E S U L T S Based on the data we collated, the Australian flowering vascular flora comprises 222 families, 2638 genera and 18,821 species of which c. 10.6% of species are introduced (Table 1). The geometric mean number of species in a family is 13.55 ± 17.08 (median ¼ 11.5, range 1–2509), and the geo- metric mean of genera in a family is 4.08 ± 1.94 (median ¼ 3, range 1–256). We consider that there are only 31 flowering plant species in Australia that can be unambiguously consid- ered extinct at the present time (Table 1). The number of families containing endangered species is 63 (179 genera), and there are 18 families (27 genera) with extinct species in them. Most of the species are herbs and shrubs(Fig. 1a), and most (c. 76%) of the introduced species in Australia are herbs (Fig. 1a). The majority of species are long-lived (more than 2 years), and introduced species are found equally in both categories (Fig. 1b). Most species have a bisexual sex system (Fig. 1c), including introduced species. The predominant fruit type is dry-dehiscent (Fig. 1d) in both native and introduced species.
Life-history characters A comparison of endangered versus extant (native species only) showed no significant difference in habit (G ¼ 6.13,
P > 0.05, Fig. 2a). However, when habit in extinct species is compared with that in extant species, trees are significantly absent from the extinct flora (G ¼ 12.76, P < 0.05, Fig. 2a). The GLM model for extinct versus extant yielded the model y habit (Wald ¼ 19.86,
P ¼ 0.013,
32% deviance
explained). A comparison of endangered versus extant for sex
system showed
significant differences (G ¼ 27.43,
P < 0.001, Fig. 2b), but not in the extinct flora (G ¼ 0.38,
P > 0.05, Fig. 2b). Similarly there were significant differences in the distribution of fruit types between endangered versus extant flora (G ¼ 56.47, P < 0.001, Fig. 2c), but not for the extinct species (G ¼ 0.25, P > 0.05, Fig. 2c). Stepwise reduction of the saturated GLM model for life history of endangered versus extant (native only) yielded y habit + sex + fruit + habit : sex + habit : fruit. We at- tempted to reduce the model further by removing the two-way interactions. An analysis of deviance indicated that removal of either of these interactions resulted in significant increases in deviance. Hence these two two-way interactions were kept in the model. Because neither the contrast for lianes nor for vines was significant when compared with other habits, we attempted to reduce the number of levels for habit by lumping lianes with shrub, and vines with herbs to fit a simpler model. Because the simpler model had a decreased AIC value and an analysis of deviance indicated that lumping habit levels did not show a significant increase in deviance, the simpler model was accepted. Further attempts to reduce the model by either removing terms or reducing the number of levels within variables either increased the AIC, resulted in a significant increase in deviance, or were not biologically relevant. Thus, the minimum model accepted is y habit3 + sex + fruit + habit3 : sex + habit3 : fruit, where habit3 consists of herbs (including vines), shrubs (including lianes) and trees (Table 2). The abundance of sex and fruit types varied significantly between the endangered and the extant data sets (Table 3). Significant interactions occurred where habit and sex type and habit and fruit type varied differently in the endangered data set compared with in the Life-history characters in the Australian flora Journal of Biogeography 33, 271–290 275
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd Table 1 A list of the families, number of genera and species and the percentage of species introduced to Australia. The endangered species from the EPBC Act (1999) are listed. Where more than one infra-specific taxon is listed only one was included in the analysis. Extinct species are listed according to rules listed in the text. Family names follow Stevens (2001 onwards) Family
No. genera No. spp.
% intro Endangered Extinct Acanthaceae 25 57 17.5 Graptophyllum reticulatum, Isoglossa eranthemoides, Xerothamnella herbacea Actinidaceae 1 1 0.0 Agavaceae 6 9
Aizoaceae 19 61 36.1 Akaniaceae 1 1
Alismataceae 5 10 60.0 Alseuosmiaceae 2 2
Amaranthaceae 47 481 8.7 Roycea pycnophylloides, Sclerolaena napiformis Anacardiaceae 10 14 35.7 Anarthriaceae 2 7
Annonaceae 19 50 4.0 Apiaceae
41 185
15.1 Eryngium fontanum, Gingidia montana, Trachymene saniculifolia Apocynaceae 44 183
9.3 Cynanchum elegans, Ochrosia moorei, Parsonsia dorrigoensis, Tylophora linearis, T. rupicola, T. woollsii Marsdenia araujacea Aponogetanaceae 1 10
Aponogeton bullusus, A. proliferus Aquifoliaceae 2 5
Araceae 16 39 15.4 Araliaceae 11 46
Astrotricha roddii Arecaceae 24 59
Archontophoenix myolensis, Ptychosperma bleeseri Aristolochiaceae 2 13
Asphodelaceae 2 5 100.0 Asteraceae 256 1121
21.9 Argentipallium spiceri, Brachyscome muelleri, Calotis moorei, Leucochrysum albicans var. tricolor, Olearia flocktoniae, Olearia hygrophila, Olearia microdisca, Rutidosis leptorrhynchoides, Senecio behrianus Olearia oliganthema, Ozothamnus selaginoides, Picris compacta, Picris drummondii, Senecio georgianus Atherspermaceae 1 1
Austrobaileyaceae 1 2 0.0 Balanopaceae 1 1
Balanophoraceae 1 2 0.0 Balsaminaceae 1 1
Basellaceae 1 1 100.0 Bataceae
1 1 0.0 Berberidaceae 2 3 100.0 Betulaceae 1 1
Bignoniaceae 12 19 36.8 Bixaceae
2 4 25.0 Blandfordiaceae 1 4 0.0 Boraginaceae 26 156
19.9 Brassicaceae 60 195
37.9 Ballantinia antipoda, Barbarea australis, Irenepharsus trypherus, Lepidium hyssopifolium, L. monoplocoides, L. peregrinum Lepidium drummondii Bromeliaceae 2 2 100.0 Burmanniaceae 2 4
Burmannia sp. Melville Island (R. Fensham 1021) Burseraceae 2 5 0.0 Byblidaceae 1 2
Cabombaceae 2 2 50.0 Cactaceae 7 23
Calycanthaceae 1 1 0.0 Campanulaceae 9 64
Hypsela sessiliflora A. Sjo¨stro¨m and C. L. Gross 276 Journal of Biogeography 33, 271–290 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd Table 1 continued Family
No. genera No. spp.
% intro Endangered Extinct Cannabaceae 2 2 100.0 Cannaceae 1 1 100.0 Caprifoliaceae 4 8
Cardiopteridaceae 1 1 0.0 Caricaceae 1 1
Caryophyllaceae 24 83 62.7 Sagina diemensis Casuarinaceae 3 64 0.0 Allocasuarina defungens, A. emuina, A. glareicola, A. portuensis, A. thalassoscopica Celastraceae 19 57
Apatophyllum constablei Centrolepidaceae 3 31
Centrolepis caespitosa Cephalotaceae 1 1
Ceratophyllaceae 1 2 0.0 Chrysobalanaceae 2 2
Cistaceae 1 1 100.0 Clusiaceae 6 23
Colchicaceae 8 47 4.3 Wurmbea calcicola, W. tubulosa Combretaceae 6 36 2.8 Commelinaceae 11 35
Connaraceae 2 2 0.0 Convolvulaceae 20 119
16.8 Cornaceae 1 2
Corsiaceae 1 1 0.0 Corynocarpaceae 1 2
Costaceae 2 3 0.0 Crassulaceae 6 29
Cucurbitaceae 18 43 27.9 Mukia sp. Longreach (D.Davidson AQ279935) Cunoniaceae 16 35 0.0 Davidsonia jerseyana, D. johnsonii Cymodoceaceae 5 10 0.0 Cyperaceae 45 682
6.6 Cyperus cephalotes, Fimbristylis adjuncta, Lepidosperma rostratum Datiscaceae 1 1
Dichapetalaceae 1 1 0.0 Dilleniaceae 5 140
0.0 Dioscoreaceae 2 7
Dipsacaeae 2 4 100.0 Doryanthaceae 1 2
Droseraceae 2 55 0.0 Ebenaceae 1 15
Diospyros mabacea Elaeagnaceae 1 1
Elaeocarpaceae 9 88 0.0 Elaeocarpus sp. Rocky Creek (G.Read AQ562114), E. williamsianus, Tetratheca deltoidea, T. gunnii, T. paynterae Tetratheca fasciculata Elatinaceae 2 8
Ericaceae 35 433 1.8 Andersonia axilliflora, A. gracilis, Epacris acuminate, E. apsleyensis, E. barbata, E. exserta, E. grandis, E. hamiltonii, E. limbata, E. sp. aff. virgata ‘graniticola’, E. stuartii, Leucopogon confertus, L. gnaphalioides, L. marginatus, L. obtectus, L. sp. Coolmunda (D.Halford Q 1635), Melichrus hirsutus J.B.Williams ms., M. sp. Gibberagee (A.S.Benwell & J.B.Williams 97239) Leucopogon cryptanthus Eriocaulaceae 1 26
Eriocaulon australasicum, E. carsonii Erythroxylaceae 1 4
Euphorbiaceae 52 278 12.2 Bertya ingramii, B. sp. Beeron Holding (P.I.Forster 5753), Beyeria lepidopetala, Fontainea oraria, Ricinocarpos trichophorus Sankowskya stipularis Amperea xiphoclada var. pedicillata Eupomatiaceae 1 2
Life-history characters in the Australian flora Journal of Biogeography 33, 271–290 277 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd Table 1 continued Family
No. genera No. spp.
% intro Endangered Extinct Fabaceae
176 2509
7.7 Acacia aprica, A. aristulata, A. ataxiphylla subsp. magna, A. auratiflora, A. brachypoda, A. cochlocarpa subsp. cochlocarpa, A. cretacea, A. enterocarpa, A. gordonii, A. insolita subsp. recurva, A. lanuginophylla, A. leptalea, A. lobulata, A. pharangites, A. pinguifolia, A. porcata, A. pygmaea, A. recurvata, A. rhamphophylla, A. ruppii, A. sciophanes, A. shirleyi, A. sp. Dandaragan (S. van Leeuwen 269), A. subflexuosa subsp. capillata, A. terminalis subsp. terminalis, A. vassalii, A. volubilis, A. whibleyana, Cajanus mareebensis, Chorizema humile, Chorizema varium, Cullen parvum, Daviesia bursarioides D. cunderdin, D. euphorbioides, D. megacalyx, D. microcarpa, D. speciosa, Gastrolobium glaucum, G. graniticum, G. hamulosum, Jacksonia pungens J.Chappill ms., J. quairading J.Chappill ms., J. sp. Collie (C.J.Koch 177), Kennedia macrophylla, Pultenaea parris parrisiae subsp. elusa, Stonesiella (Pultenaea) selaginoides, Swainsona recta Acacia kingiana, A. prismifolia, Indigofera efoliata, Pultenaea maidenii, Streblorrhiza speciosa Fagaceae
1 3 100.00 Flagellariaceae 1 1 0.0 Frankeniaceae 1 47
Frankenia plicata Fumariaceae 4 10
Gentianaceae 8 31 22.6 Gentiana baeuerlenii, G. wingecarribiensis Geraniaceae 3 34 41.2 Gesneriaceae 5 6
Goodeniaceae 12 379 0.0 Coopernookia georgei, Lechenaultia laricina, L. pulvinaris Grossulariaceae 10 27 7.4 Gunneraceae 1 1
Gyrostemnonaceae 5 18 0.0 Gyrostemon reticulatus Haemodoraceae 8 74 2.7 Anigozanthos bicolor subsp. minor, Conostylis dielsii subsp. teres, C. drummondii, C. lepidospermoides, C. micrantha, C. misera, C. seorsiflora subsp. trichophylla, C. setigera subsp. dasys, C. wonganensis Haloragaceae 6 105
0.0 Haloragis eyreana, Haloragodendron lucasii, Myriophyllum lapidicola Haloragis platycarpa Hamamelidaceae 3 3 0.0 Hanguanaceae 1 1
Hemerocallidaceae 1 1 0.0 Hernandiaceae 3 6
Himantandraceae 1 1 0.0 Hydatellaceae 2 7
Hydatella dioica Hydrocharitaceae 13 28
Hypoxidaceae 1 10 0 Icacinaceae 6 7
Iridaceae 32 95 69.5 Orthrosanthus muelleri, Patersonia spirafolia Juncaceae 2 74 27.0 Juncaginaceae 3 16
Lamiaceae 44 293 16.0 Hemiandra gardneri, H. rutilans, H. sp. Watheroo (S.Hancocks 4), Plectranthus habrophyllus, P. nitidus, P. omissus, P. torrenticola rostanthera askania, P. eurybioides, P. junonis, Westringia crassifolia, W. kydrensis Prostanthera albo-hirta, P. marifolia Lauraceae 9 131
1.5 Endiandra cooperana, E. floydii Laxmanniaceae 3 107 0.0 Lecythidaceae 2 5
Lentibulaniaceae 1 61 1.6 Liliaceae 55 180
24.4 Borya mirabilis, Dianella amoena Limnocharitaceae 2 2 50.0 A. Sjo¨stro¨m and C. L. Gross 278 Journal of Biogeography 33, 271–290 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd Table 1 continued Family
No. genera No. spp.
% intro Endangered Extinct Linaceae
1 7 71.4 Loganiaceae 9 101 1.0 Loranthaceae 12 69
Amyema scandens Luzuriagaceae 1 2
Lythraceae 10 29 17.2 Malpighiaceae 2 2
Malvaceae 61 471 7.0 Corchorus cunninghamii, Lasiopetalum pterocarpum E.M.Benn & K.Shepherd ms., L. rotundifolium, Rulingia prostrata, Rulingia sp. Trigwell Bridge (R.Smith s.n. 20/6/1989), Thomasia sp. Green Hill (S.Paust 1322) Melastomataceae 5 8
Meliaceae 12 41 2.4 Melianthaceae 1 2
Memecylaceae 1 3 0.0 Menispermaceae 11 18
Carronia pedicellata Menyanthaceae 3 30
Villarsia calthifolia Molluginaceae 3 13
Macarthuria keigheryi Monimiaceae 12 35
Daphnandra sp. Illawarra (R.Schodde 3475) Moraceae
10 46 4.3 Moringaceae 1 1 100.0 Musaceae
1 3 0.0 Musa fitzalanii Myoporaceae 2 248
0.0 Eremophila denticulata subsp. trisulcata Chinnock ms., E. nivea, E. pinnatifida Chinnock ms., E. resinosa, E. scaberula, E. ternifolia, E. veneta Chinnock ms., E. verticillata, E. virens, Myoporum turbinatum Myristicaceae 2 3
Myrsinaceae 7 34 5.9 Rapanea sp. Richmond River (J.H.Maiden & J.L.Boorman NSW 26751) Myrtaceae 77 1593 0.4 Austromyrtus fragrantissima, Austromyrtus gonoclada, Baeckea kandos, Calytrix breviseta subsp. breviseta, Chamelaucium sp. Gingin (N.G.Marchant s.n. 4/11/1988), Darwinia acerosa, D. apiculata, D. carnea, D. chapmaniana Marchant & Keighery ms., D. collina, D. ferricola N.G.Marchant & Keighery ms., D. oxylepis, D. sp. Carnamah (J.Coleby-Williams 148), D. sp. Williamson (G.J.Keighery 12717), D. wittwerorum, Decaspermum sp. Mt Morgan (D.Hoy 71), Eucalyptus absita, E. balanites, E. beardiana, E. bennettiae, E. brevipes, E. burdettiana, E. conglomerata, E. copulans, E. crenulata, E. crucis subsp. praecipua, E. cuprea, E. dolorosa, E. graniticola Brooker & Hopper ms., E. gunnii subsp. divaricata, E. imlayensis, E. impensa, E. insularis, E. leprophloia, E. morrisbyi, E. pachycalyx subsp. banyabba, E. phylacis, E. pruiniramis, E. recurva, E. rhodantha var. petiolaris, E. sp. Howes Swamp Creek (M.Doherty 19/7/1985 NSW 207054), Hypocalymma longifolium, Micromyrtus grandis, Triplarina imbricata, Triplarina nowraensis, Uromyrtus australis, Verticordia albida, V. densiflora var. pedunculata, V. fimbrilepis subsp. fimbrilepis,V. harveyi,V. hughanii, V. pityrhops, V. plumosa var. ananeotes, V. plumosa var. pleiobotrya, V. plumosa var. vassensis, V. spicata subsp. squamosa, V. staminosa subsp. cylindracea var. cylindracea, V. staminosa subsp. staminosa, Xanthostemon formosus Nelumbonaceae 1 1
Nepenthaceae 1 1 0.0 Nothofagaceae 1 3
Nyctaginaceae 5 15 13.3 Nymphaeceae 2 13
Ochnaceae 2 2 50.0 Olacaceae 3 13
Oleaceae 9 30 20.0 Life-history characters in the Australian flora Journal of Biogeography 33, 271–290 279
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd Table 1 continued Family
No. genera No. spp.
% intro Endangered Extinct Onagraceae 5 37 54.1 Opiliaceae 2 2 0.0 Orchidaceae 105 862
0.3 Arthrochilus huntianus subsp. nothofagicola, Caladenia (listed as Drakonorchis) drakeoides Hopper & A.P.Brown ms., C. amoena, C. anthracina, C. arenaria, C. argocalla, C. atroclavia, C. audasii, C. behrii, C. bryceana subsp. bryceana, C. busselliana Hopper & A.P.Brown ms., C. caesarea subsp. maritima, C. campbellii, C. carnea var. subulata, C. colorata, C. dienema, C. dorrienii, C. elegans Hopper & A.P.Brown ms., C. excelsa Hopper & A.P.Brown ms., C. fragrantissima subsp. orientalis, C. fulva, C. gladiolata, C. hastata, C. hoffmanii Hopper & A.P.Brown ms., C. huegelii Hopper & A.P.Brown ms., C. lindleyana, C. lowanensis, C. macroclavia, C. pallida, C. richardsiorum, C. rigida, C. robinsonii, C. rosella, C. saggicola, C. sp. aff. venusta, C. sylvicola, C. tensa, C. thysanochila, C. tonellii, C. viridescens Hopper & A.P.Brown ms., C. winfieldii Hopper & A.P.Brown ms., C. xanthochila, C. xantholeuca, Calochilus psednus, C. richiae, Corybas sp. Finniss (R.Bates 28794), Crepidium lawleri, Dendrobium antennatum, D. lithocola, D. mirbelianum, D. nindii, Dipodium pictum, Diuris basaltica D.L. Jones ined., D. fragrantissima, D. lanceolata, D. pedunculata, D. purdiei, Drakaea confluens Hopper & A.P.Brown ms., D. elastica, D. isolata Hopper & A.P.Brown ms., Epiblema grandiflorum var. cyanea K.Dixon ms., Genoplesium brachystachyum, G. firthii, G. plumosum, G. rhyoliticum, G. tectum, Habenaria macraithii, Microtis angusii, Paracaleana dixonii Hopper & A.P.Brown ms., Phaius australis, P. bernaysii, Phaius tancarvilleae, Phalaenopsis rosenstromii, Prasophyllum affine, P. amoenum, P. apoxychilum, P. castaneum, P. correctum, P. diversiflorum, P. favonium, P. frenchii, P. milfordense, P. olidum, P. perangustum, P. petilum, P. pulchellum, P. robustum, P. secutum, P. stellatum, P. suaveolens, P. subbisectum, P. tunbridgense, P. uroglossum, Pterostylis aenigma, P. atriola, P. basaltica, P. commutata, P. despectans, P. gibbosa, P. rubenachii, P. saxicola, P. sp. Botany Bay (A.Bishop J221/1-13), P. sp. Halbury (R.Bates 8425), P. sp. Hale (R.Bates 21725), P. sp. Northampton (S.D.Hopper 3349), P. wapstrarum, P. ziegeleri, Rhizanthella gardneri, Thelymitra epipactoides, T. jonesii, T. manginii K.Dixon & Batty ms., T. stellata, Vrydagzynea paludosa Acianthus ledwardii, Caladenia brachyscapa, C. pumila, Diplocaulobium masonii, Diuris bracteata, Oberonia attenuata Orobanchaceae 1 2 50.0 Oxalidaceae 2 30
Pandanaceae 2 36 0.0 Pandanus spiralis var. flammeus Papaveraceae 6 13 100.0 Passifloraceae 2 14
Pedaliaceae 5 8 50.0 Philydraceae 3 5
Phyllanthaceae 14 148 2.0 Phytolaccaceae 3 5
Piperaceae 3 11 9.1 Pittosporaceae 11 88
Bentleya spinescens, Marianthus (Billardiera) mollis Plantaginaceae 2 43
Plumbaginaceae 4 11 63.6 Poaceae
234 1299
23.9 Agrostis adamsonii, A. limitanea, A. granitica, Austrostipa wakoolica, Danthonia popinensis, Deyeuxia appressa, D. drummondii, Digitaria porrecta, Eremochloa muricata, Glyceria drummondii Amphibromus whitei
Podostemaceae 2 2 0.0 Polemoniaceae 3 4
Polygalaceae 6 45 11.1 Polygonaceae 11 54
Muehlenbeckia tuggeranong Pontederiaceae 3 6
Portulacaceae 9 59 8.5 A. Sjo¨stro¨m and C. L. Gross 280 Journal of Biogeography 33, 271–290 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd Table 1 continued Family
No. genera No. spp.
% intro Endangered Extinct Posidonaceae 1 8 0.0 Potamogetonaceae 4 19 5.3 Primulaceae 4 10
Proteaceae 46 1007 0.0 Adenanthos cunninghamii, A. dobagii, A. eyrei, A. pungens subsp. effusus, A. velutinus, Banksia brownii, B. cuneata, B. oligantha, Conospermum densiflorum subsp. unicephalatum, Conospermum toddii, Dryandra anatona, D. aurantia, D. ionthocarpa, D. mimica, D. montana, D. nivea subsp. uliginosa, Eidothea hardeniana, Grevillea acanthifolia subsp. paludosa, G. althoferorum, G. batrachioides, G. beadleana, G. caleyi, G. calliantha, G. christineae, G. curviloba subsp. curviloba, G. curviloba subsp. incurva, G. dryandroides subsp. dryandroides, G. dryandroides subsp. hirsuta, G. guthrieana, G. humifusa, G. iaspicula, G. infundibularis, G. involucrata, G. maccutcheonii, G. masonii, G. maxwellii, G. mollis, G. molyneuxii, G. murex, G. obtusiflora, G. pythara, G. rara, G. rivularis, G. scapigera, G. wilkinsonii, G. williamsonii, Hakea dohertyi, H. pulvinifera, Isopogon uncinatus, Lambertia echinata subsp. echinata, L. echinata subsp. occidentalis, L. fairallii, L. orbifolia, Lomatia tasmanica, Macadamia jansenii, Persoonia hirsuta, P. micranthera, P. mollis subsp. maxima, P. nutans, Petrophile latericola Keighery ms., Synaphea quartzitica, Triunia robusta Persoonia laxa,
P. prostrata Putranjivaceae 1 4
Rafflesiaceae 1 2 0.0 Ranunculaceae 9 67
Ranunculus prasinus Resedaceae 1 4
Restionaceae 35 119 0.0 Chordifex (Restio) abortivus Rhamnaceae 18 165 1.8 Pomaderris cotoneaster, Spyridium microphyllum, Spyridium sp. (Little Desert), Stenanthemum pimeleoides Rhipogonaceae 1 5
Rhizophoraceae 4 12 0.0 Rosaceae
22 82 76.8 Rubiaceae 49 248 6.9 Randia moorei Rutaceae 46 397 1.3 Acronychia littoralis, Asterolasia elegans, Boronia capitata subsp. capitata, B. exilis, B. granitica, B. repanda, B. revoluta, Correa lawrenceana var. genoensis, Drummondita ericoides, Leionema equestre, L. lachnaeoides, Phebalium daviesii, Philotheca basistyla, P. freyciana, P. wonganensis, Zieria adenophora, Z. baeuerlenii, J.A.Armstrong ms., Z. buxijugum J.Briggs & J.A.Armstrong ms., Z. covenyi J.A.Armstrong ms., Z. floydii J.A.Armstrong ms., Z. formosa J.Briggs & J.A.Armstrong ms., Z. granulata, Z. ingramii J.A.Armstrong ms., Z. lasiocaulis J.A.Armstrong ms., Z. obcordata, Z. parrisiae J.Briggs & J.A.Armstrong ms., Z. prostrata J.A.Armstrong ms., Z. sp. Brolga Park (A.R.Bean 1002) Philotheca falcatus Salicaceae 11 31
Santalaceae 13 61 0.0 Spirogardnera rubescens Sapindaceae 33 200 2.5 Alectryon ramiflorus, Atalaya collina, Cossinia australiana, Diploglottis campbellii, Dodonaea subglandulifera, Toechima pterocarpum Sapotaceae 7 29
Pouteria (Planchonella) eerwah Saxifragaceae 1 1
Scrophulariaceae 51 170 30.0 Euphrasia collina subsp. muelleri, E. collina subsp. osbornii, E. fragosa, E. gibbsiae subsp. psilantherea, E. semipicta, E. sp. ‘fabula’, Microcarpaea agonis, Stemodia haegii Euphrasia arguta
Simaroubaceae 4 9 11.1 Quassia sp. Mooney Creek (J.King s.n. 1949) Smilaceae 3 9 0.0 Solanaceae 23 210
24.3 Cyphanthera odgersii subsp. occidentalis, Symonanthus bancroftii Sparganiaceae 1 2 50.0 Sphenocleaceae 1 1
Stemonaceae 1 4 0.0 Life-history characters in the Australian flora Journal of Biogeography 33, 271–290 281
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd extant data set (Fig. 3, Table 3), notably in the herbaceous life form.
Stepwise reduction of the multinomial model comparing endangered, extinct and extant species yielded y habit + sex + fruit + habit : fruit. As for the GLM model for the endangered species, the model was further reduced by lumping the habit categories of herbs with vines, and shrubs with lianes. This simpler model had a reduced AIC, and an anova comparing the two models showed no significant increase in deviance (LR ¼ 12.66, d.f. ¼ 12, P > 0.05). Species richness When the abundance of endangered species was examined across the 1978 native genera, most of them contained no endangered species. The GLM regression of proportion of endangered to the log of the size of genus was highly significant (odds ratio ¼ 1.131 with 95% confidence interval (CI) 1.099– 1.163, t
¼ 4.37, P < 0.001; G 2 ¼ 19.17, d.f. ¼ 1, P < 0.001, Fig. 4a), showing that species-rich genera are proportionately more likely to contain endangered species than species-poor genera. These results are not surprising: the bigger the genus the more likely it will be that it will contain an endangered species. When only those genera containing an endangered species were examined (n ¼ 179), endangered species are more likely to occur in small genera (odds ratio ¼ 0.619 with 95% CI 0.596–0.643, t ¼ )7.17; G
2 ¼ 169.53, d.f. ¼ 1, P < 0.001, Fig. 4c). However, because most genera are small in the Australian flora we queried whether this relationship would hold irrespective of which genera are used in the analyses. If a significant relationship between genus size and the proportion of endangered species can be detected using a random sample of all genera it would indicate that endangered species are occurring in small genera in a random way. This was tested using a bootstrap test on a model with the original endangered proportion data as the response variable and the log of the size of the genus as the explanatory term. A weighted regression was performed on each of 10 000 different random samples (n ¼ 179). Of the resulting P values, only 2945 were ‡ 0.05. This strongly suggests that endangered species only occur in small genera because most genera in Australia are small. Most genera do not have extinct species, but no significant relationship was found against genus richness (t ¼ )0.18;
G 2 ¼ 0.61, d.f. ¼ 1, P ¼ 0.43, Fig. 4b). When only those genera containing an extinct species were examined (n ¼ 27),
a significant relationship was found whereby small genera are more likely to contain extinct species than larger genera (odds ratio ¼ 0.365
with 95%
CI 0.318–0.420, t ¼
G 2 ¼ )61.54, d.f. ¼ 1, P < 0.001, Fig. 4d). Only 33% of families contained at least one endangered or extinct species. Similarly, the relationship between threat status and family speciosity showed that highly species-rich families are more likely to contain endangered species than species-poor families (odds ratio ¼ 1.139 with 95% CI 1.073–1.209, t ¼ 2.18; G 2 ¼ 14.5, d.f. ¼ 1, P ¼ 0.03). A significant rela- tionship between species richness in families and extinct species was not found (t ¼ 0.51; G 2 ¼ 0.26, d.f. ¼ 1, P ¼ 0.61). When only those families containing an endangered species were Table 1 continued Family No. genera No. spp. % intro
Endangered Extinct
Stylidiaceae 5 241 0.0 Stylidium coroniforme Surianaceae 4 5 0.0 Symplocaceae 1 15
Tamaricaceae 1 1 100.0 Theaceae
2 2 50.0 Thymelaceae 10 106 0.9 Pimelea spicata, P. spinescens subsp. spinescens, P. venosa Pimelea spinescens subsp. pubiflora Trimeniaceae 1 1
Triuridaceae 1 1 0.0 Tropaeolaceae 1 1
Typhaceae 1 3 33.3 Ulmaceae
4 10 10.0 Urticaceae 13 25 28.0 Valerianaceae 3 8
Verbenaceae 28 167 10.8 Pityrodia scabra Violaceae 5 23 13.0 Vitaceae
7 44 2.3 Winteraceae 2 10 0.0 Xanthorrhoeaceae 9 50
Xyridaceae 1 20 0.0 Zingiberaceae 9 18
Zosteraceae 2 4 0.0 Zygophyllaceae 5 55
Total 2638
18,821 10.60
A. Sjo¨stro¨m and C. L. Gross 282
Journal of Biogeography 33, 271–290 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd used in the analyses (n ¼ 62), no significant relationship was detected (t ¼ )0.68; G 2 ¼ 3.24, d.f. ¼ 1, P ¼ 0.5). However, when only those families containing an extinct species were used in the analyses, a significant relationship was detected (odds ratio ¼ 0.538 with 95% CI 0.465–0.621, t ¼ )4.23; G
¼ 14.98, d.f. ¼ 1, P < 0.0001), showing that large families are less likely to have extinct species than smaller families. However, to examine this relationship further for family size and families containing only extinct species we ran another bootstrap test (see above). Here the results from the weighted regressions using 10 000 different random samples of families (n ¼ 16) and the proportion of extinct species were mostly significant, with 4144 of the 10 000 iterations yielding a P value ‡ 0.05. Thus low family speciosity is an unlikely predictor of extinction risk. The association of extinction risk with phylogeny The distribution of threat status at the generic level was investigated for 1640 genera in the Australian angiosperm 0 10
30 40 50 Herb Vine Liane Shr ub Tree
% of species % of species All
Extinct Endangered 0 20
60 80 100 Bisexual Dioecious Monoecious 0 20 40 60 80 Dry-indeh Dry-deh
Fleshy Kataloq: teaching teaching -> Oncological Emergencies Dr. Gary Harding md, frcpc teaching -> Chapter S: IV iv. Informed Search teaching -> Exploring a Protein Structure in the rcsb pdb: Insulin Learning Goals teaching -> Infertility, In Vitro Fertilization (ivf) and Genetic Testing Susan Sarajari, M. D., Ph. D teaching -> Java – gui graphical User Interface ine2720 Web Application Software Development teaching -> Exploring a Protein Structure in the rcsb pdb: Antibodies Learning Goals teaching -> Malrotation and volvulus teaching -> Pathogenic Bacteria/4 3rd Lab. Investigation Pathogenesis a microorganism is a pathogen teaching -> Another length Page of 1 1 metre = 1000 millimetres teaching -> Iteration 1 First real development iteration Yüklə 225,04 Kb. Dostları ilə paylaş:
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