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— Acanthaceae is a family of tropical flowering plants with approximately 4900 species. Despite remarkable variation in morphological traits, research on patterns of character evolution has been limited by uncertain relationships among some of the major lineages. We sampled 16 taxa from these major lineages to estimate a phylogenomic framework using a combination of five newly sequenced shotgun genome skims plus seven new and four publicly available transcriptomes. We used OrthoFinder2 to infer a species tree with strong branch support. Except for the placement of Crabbea, our results corroborate the most recent chloroplast and nrITS sequence-based topology. Of 587 single copy loci, 10 were recovered for all 16 species; a RAxML tree estimated from these 10 loci resulted in the same topology as other datasets assembled in this study, with the exception of relationships among three sampled species of Barleria; however, branch support was lower compared to the tree reconstructed using more data. ABBA-BABA tests were conducted to investigate patterns of introgression involving Crabbea; few nucleotides supported alternative topologies. SplitsTree networks of the 587 loci and 6136 orthogroup trees revealed conflict among the branches leading to Andrographideae, Whitfieldieae, and Neuracanthus. A principal components analysis in treespace found no distinct clusters of trees. Our results based on combined genome skim and transcriptome sequences strongly corroborate the previously published chloroplast and nr-ITS-based phylogeny of Acanthaceae with increased resolution among Barlerieae, Andrographideae, Whitfieldieae, and Neuracanthus. This advance in our knowledge of Acanthaceae relationships will allow us to investigate character evolution and other phenomena within this diverse group of plants in studies with increased taxon sampling.

We present phylogenetic results for Justicieae, the largest of the major lineages of Acanthaceae. With 300% more sequence data and a similarly increased but also more geographically representative taxon sample compared to the only previous study of the lineage as a whole, we confirm the existence of the Pseuderanthemum Lineage, Isoglossinae, the Tetramerium Lineage and the Justicioid Lineage. To the last three lineages, we add a number of taxa with the goal of advancing our knowledge of genera (e.g., Isoglossa, Rhinacanthus) and of geographic areas (e.g., Malagasy and Asian Acanthaceae). These added taxa are accommodated within the phylogenetic framework for the lineages established in our earlier work with one exception. The monospecific genus Ichthyostoma is not placed in any of the established lineages but is instead sister to (Tetramerium Lineage + Justicioid Lineage). As this result is unexpected, we examined topologies both including and excluding this plant which revealed that other aspects of relationships remain stable whether Ichthyostoma is included or not. We point to Ichthyostoma as one of several Justicieae that will benefit from additional study. One species of Isoglossa is placed in the Tetramerium Lineage instead of with all other sampled members of the genus in Isoglossinae. Pollen morphology of I. variegata is more consistent with its phylogenetic placement here than with its taxonomic assignment to Isoglossa. As suggested in our earlier work, pollen morphology can be a powerful signal of phylogenetic relationships in Justicieae. With our increased sampling, we show that the Ptyssiglottis Lineage, a relatively small group of African and Asian plants, warrants recognition separate from Isoglossinae. With the addition of the Ptyssiglottis Lineage and excluding Ichthyostoma for now, the lineages of Justicieae are related as follows: (Pseuderanthemum Lineage ((Isoglossinae + Ptyssiglottis Lineage) (Tetramerium Lineage + Justicioid Lineage))). The Pseuderanthemum Lineage has been understudied to date and we here attempt to include representatives of all genera that are putatively part of this lineage and also to test monophyly of the larger genera. We identify a number of strongly supported clades but some aspects of relationship remain unresolved. Asystasia, Graptophyllum, Oplonia, and Pseuderanthemum are not monophyletic although, with a small number of nomenclatural changes, the first and second can be rendered monophyletic. With a range essentially as extensive as that of Justicieae as a whole, the Pseuderanthemum Lineage is biogeographically complex with patterns of distribution suggestive of multiple shifts between continents including Australia and parts of the Pacific basin. The calibrated phylogeny confirms that intercontinental shifts in distribution across Justicieae must have been by long-distance dispersal rather than by vicariance or stepwise dispersal over land bridges and also points to a number of groups that have potentially undergone rapid diversification. The Pseuderanthemum Lineage, in particular, requires considerable additional research to both understand relationships and achieve an informative taxonomy for the group. Although we here establish a phylogenetic framework across Justicieae, Next Generation Sequence data will be necessary to elucidate details of relationships in most lineages. Additional study of structural characters is also warranted as we continue to be unable to identify structural synapomorphies for a number of aspects of phylogenetic relationships that are very strongly supported by molecular data.

Increases in the incidence and severity of drought threaten the viability of rare plants in arid regions. The endangered Nichol's Turk's head cactus (Echinocactus horizonthalonius Lemaire var. nicholii L. Benson) occurs only in four small, isolated populations in the Sonoran Desert of North America. Since 1995 we have monitored a population in southeastern Arizona (USA). Here we report 23 years of observations on abundance, growth, mortality, flowering and recruitment. Abundance of plants decreased from 132 in 1996 to 40 in 2017, with 100 individuals recruited and 203 dying during the study. Individual plants grew slowly, increasing annually by an average of 0.22 cm (95% confidence interval, 0.18–0.26 cm) in diameter and 0.27 cm (0.20–0.33 cm) in height. Growth was slowest when drought was most severe and slowed as plants reached maximum size. Annual mortality increased markedly across the study period and did not vary with plant size. Annual probability of flowering increased as plants increased in diameter but not in height, and varied with precipitation and drought but not with mean annual temperature. Recruitment was higher when average temperature was higher and the number of recruits per capita increased across the study period. The annual rate of change in abundance averaged −6%, but shifted markedly from −1% during 1995–2008 to −11% during 2008–2017. Our results indicate that the population's decline was not a consequence of failed recruitment but of increased mortality, which we discuss in the context of climate and herbivory.

The New World ‘justicioid’ lineage (Acanthaceae: Acanthoideae: Justicieae) includes about 500 species and encompasses extensive variation in flower shape, size, and color along with remarkable diversity in anther, seed, and pollen mophology. Species richness of this lineage is due to the inclusion of the New World members of the polyphyletic genus Justicia in addition to five smaller Neotropical genera: Cephalacanthus, Clistax, Harpochilus, Megaskepasma, and Poikilacanthus. Although previous studies have shown that the New World ‘justicioid’ lineage is monophyletic, none have focused specifically on relationships among these plants nor has the current infrageneric classification of Justicia for the New World been tested. This is the first phylogenetic study to include a dense sample of taxa from the New World ‘justicioid’ lineage. We use DNA sequence data for five plastid, nuclear ribosomal ITS, and one low copy nuclear region (8315 bp) for 134 in-group taxa to estimate the phylogenetic framework for the group, and test the current infrageneric classification of New World Justicia. Most of the currently recognized sections of New World ‘justicioids’ are not monophyletic. On the other hand, our phylogenetic results identify nine major clades, many of which are supported by a combination of structural traits. The results of this study provide a foundation for the placement of other species of New World Justicia that have yet to be sampled for phylogenetic analysis, as well as for future comparative studies of evolutionary processes and revisionary work on this diverse and complex lineage.

Molecular data confirm monophyly of the Tetramerium lineage (Acanthaceae, Justicieae), a group of nearly 200 species, ca. 75% from the New World (NW) and the remainder from Asia or Africa. Compared to our earlier work, . 50% more in-group taxa were sampled (nearly 2/3 of known taxa now sampled) and added ∼20% more sequence data. We time-calibrate the phylogeny using primary fossil evidence and the larger context of a calibrated phylogeny for Acanthaceae. Many aspects of relationship are strongly supported but uncertainty remains about a number of phylogenetically isolated taxa (e.g. African Angkalanthus, Chorisochora; NWAncistranthus) and some aspects of relationships among clades remain weakly supported. The group originated in the Old World (OW) and began diversifying about 11.5 Ma. A single dispersal event to the NW occurred about 8.3 Ma, likely from Africa to southwestern North America. Anumber ofmajor clades, particularly in the NW, are less than 5Ma old and species in many clades are substantially younger. Several NW clades, even those with < 10 species, comprise species that are markedly heterogeneous in floral traits likely reflecting selection by pollinators. Many NW genera are not monophyletic, which is not surprising given that floral traits have been relied upon taxonomically. Diversification rate analysis revealed no significant shifts resulting in a gradual increase in number of lineages with time. The Tetramerium lineage is now phylogenetically well characterized but remains enigmatic from the perspective of morphological evidence for many aspects of relationships. The lineage is usefully thought of as a ‘clade complex’: as a species complex is of morphologically confusing species, a ‘clade complex’ is of clades that are not well characterized morphologically.

Cyrtandra (Gesneriaceae) is among the largest genera of flowering plants in the remote oceanic islands of the Pacific, with an estimated 175 species distributed across an area that extends from the Solomon Islands, east to the Marquesas Islands, and north to the Hawaiian Islands. The vast majority of species are single-island endemics that inhabit upland rainforests. Although previous molecular phylogenetic studies greatly advanced our understanding of the diversification of Pacific Cyrtandra, a number of uncertainties remain regarding phylogenetic relationships, divergence times, and biogeographic patterns within this large and widely dispersed group. In the present study, five loci (ITS, ETS, Cyrt1, psbA-trnH, and rpl32-trnL) were amplified and sequenced for phylogenetic reconstruction of 121 Cyrtandra taxa. Maximum likelihood and Bayesian inference confirmed that C. taviunensis from Fiji is sister to the remaining members of the Pacific clade. Dating analyses and ancestral area estimation indicates that the Pacific clade of Cyrtandra originated in Fiji during the Miocene ca. 9mya. All major crown lineages within the Pacific clade appeared < 5mya, coincident with the emergence of numerous Pacific islands and a subsequent increase in available habitat. The biogeographic history of Cyrtandra in the Pacific has been shaped by extinction, dispersal distance, and founder events. Biogeographic stochastic mapping analyses suggest that cladogenesis within Pacific Cyrtandra involved a combination of narrow (within-area) sympatry and founder events. A mean of 24 founder events was recovered between Pacific archipelagos, while a mean of 10 founder events was recovered within the Hawaiian archipelago.

Summary DNA sequence data, macro-morphological evidence and pollen analysis are used to clarify the phylogenetic placement of two African species of Acanthaceae: Schaueria populifolia C.B.Clarke and Rhinacanthus ndorensis Schweinf. The combined data demonstrate that these species are only distantly related to the genera in which they are currently placed and provide strong support for recognition of new genera to accommodate them. Two new genera are therefore proposed and a taxonomic account is provided for each of these. The first, Champluviera I.Darbysh., T.F.Daniel & C.A.Kiel, is based primarily on S. populifolia but Chlamydocardia nuda C.B.Clarke (= Justicia tigrina Heine) is also included within this new genus based on shared morphological traits, at least one of which is synapomorphic and restricted to these species. Based on molecular evidence, Champluviera is placed in a well-supported clade of several genera that are together sister to the core Isoglossinae lineage of tribe Justicieae. The genus may be recognised morphologically by the combination of a dense spiciform terminal thyrse with the bracts, bracteoles and calyx lobes all slender and closely resembling one another; a bilabiate corolla lacking a rugula and with intricate speckling or striping on the lips; and, in particular, the putatively synapomorphic trait of anther thecae with broad, flattened, basal appendages that have an irregularly toothed fringe. The second new genus, Kenyacanthus I.Darbysh. & C.A.Kiel, is based on R. ndorensis and is so far thought to be monospecific. It is placed between core Diclipterinae and Monechma Group II within the expanded subtribe Diclipterinae in tribe Justicieae. Kenyacanthus can be recognised morphologically by having the combination of a trailing or procumbent habit; fasciculate inflorescences; bracts, bracteoles and calyces with hyaline margins; a bilabiate corolla with a slender cylindrical tube longer than the limb and with a shallow rugula; stenotribic flowers, with the stamens held against the lower lip; and anthers with only slightly offset thecae that lack appendages.

Premise of research. Plants with C4 photosynthesis are able to produce carbohydrates more efficiently than plants with C3 photosynthesis in warm climates when levels of atmospheric CO2 are reduced. The C4 pathway has evolved multiple times in distantly related lineages, but it is not known whether the same physiological transitions occurred in all lineages. Species with intermediate C3-C4 physiology and anatomy offer the opportunity to study how plants transition from C3 to C4. It is thus vital to characterize phylogenetic relationships and photosynthetic pathways in groups with C3-C4 intermediate species, as well as C3 and C4 species.Methodology. We assessed photosynthetic pathway evolution in the Afro-Asian genus Blepharis (Acanthaceae) by sampling 99 species for carbon isotope ratios, 18 species for leaf anatomy, and 36 species for phylogenetic analysis. We estimated when Blepharis clades diverged using a BEAST molecular dating analysis, and we estimated ancestral distributions using BioGeoBEARS. We al...

The California Phenology Thematic Collections Network (CAP TCN) is a collaborative project that seeks to maximize the value of herbarium specimens and their data, especially for understanding changes in plant phenology due to anthropogenic climate change. The project unites personnel in herbaria at California universities, research stations, natural history museums, and botanic gardens with the goal of capturing images, transcribing label data, and producing georeferenced coordinates of nearly one million preserved plant specimens collected over the past 150+ years. Each digitized specimen will also be scored for its phenological status—the stage of growth and reproduction of the specimen such as flowering or fruiting. The CAP TCN is developing efficient workflows and data standards necessary to collect, store, and analyze trait data from specimens to ensure their utility for research and other applications. These novel resources and data will enable powerful research in phenology and other topics in the California Floristic Province biodiversity hotspot and beyond.

The Mirandea clade (Acanthaceae) is part of the Tetramerium lineage (Acanthaceae: Justicieae). Traditionally, macromorphological floral traits such as floral form and color have guided taxonomy in the Tetramerium lineage but earlier results on this group indicate that these traits are highly homoplastic. The present study includes all described species and multiple samples of Hoverdenia, Mirandea, and Yeatesia to fully delimit the Mirandea clade, test monophyly of species, and determine relationships among them using DNA sequence data from five regions (nuclear: nrITS and ncpGS; cp: trnS—G, ndhF—trnL(UAG), and trnT-L). The Mirandea clade is here shown to include seven species currently placed in four genera: Mirandea, Hoverdenia, Yeatesia, and Justicia. The lineage is edaphically and geographically cohesive, occurring along the Sierra Madre Oriental in Mexico and southern Texas in xeric habitats, except for one species that inhabits a broad geographical region of the southeastern U. S. A. in mesi...

Modern agricultural practices pose serious threats to biodiversity worldwide. Species losses from habitat conversion are well documented, but indirect impacts such as reduced water availability to adjacent ecosystems are less known. San Quintin is an important agricultural valley in the mediterranean climate region of Baja California, Mexico. The region is also a hotspot of plant species richness and endemism. Plant species in the region are here analyzed by comparison of the contemporary flora to historical botanical collections to identify extirpations. Historical collections indicate that habitat loss to agriculture has been a direct cause of species losses. As importantly, the unsustainable extraction of groundwater has apparently led to salt water intrusion, resulting in the loss of 22 native plant taxa, including 13 rare plants. Seventy-eight percent of all the vernal pool taxa have been lost from the flora (including 85 % of the rare taxa) and 11 % of plants of riparian and pond habitat (including 25 % of the rare taxa) are no longer found in the region. Unsustainable agricultural practices continue to threaten fragile coastal ecosystems and are a serious challenge to current and future conservation efforts. Ironically, these same practices frequently result in abandonment of cultivated areas. Owing to indirect impacts, conservation of biodiversity and large-scale agricultural operations are even less compatible on a regional scale than indicated by direct impacts. It is vital that sustainable agricultural practices be adopted locally and globally to avoid further losses of biodiversity.

More than a decade of phylogenetic research has yielded a well-sampled, strongly supported hypothesis of relationships within the large ( > 4000 species) plant family Acanthaceae. This hypothesis points to intriguing biogeographic patterns and asymmetries in sister clade diversity but, absent a time-calibrated estimate for this evolutionary history, these patterns have remained unexplored. Here, we reconstruct divergence times within Acanthaceae using fossils as calibration points and experimenting with both fossil selection and effects of invoking a maximum age prior related to the origin of Eudicots. Contrary to earlier reports of a paucity of fossils of Lamiales (an order of ∼ 23,000 species that includes Acanthaceae) and to the expectation that a largely herbaceous to soft-wooded and tropical lineage would have few fossils, we recovered 51 reports of fossil Acanthaceae. Rigorous evaluation of these for accurate identification, quality of age assessment and utility in dating yielded eight fossils judged to merit inclusion in analyses. With nearly 10 kb of DNA sequence data, we used two sets of fossils as constraints to reconstruct divergence times. We demonstrate differences in age estimates depending on fossil selection and that enforcement of maximum age priors substantially alters estimated clade ages, especially in analyses that utilize a smaller rather than larger set of fossils. Our results suggest that long-distance dispersal events explain present-day distributions better than do Gondwanan or northern land bridge hypotheses. This biogeographical conclusion is for the most part robust to alternative calibration schemes. Our data support a minimum of 13 Old World (OW) to New World (NW) dispersal events but, intriguingly, only one in the reverse direction. Eleven of these 13 were among Acanthaceae s.s., which comprises > 90% of species diversity in the family. Remarkably, if minimum age estimates approximate true history, these 11 events occurred within the last ∼ 20 myr even though Acanthaceae s.s is over 3 times as old. A simulation study confirmed that these dispersal events were significantly skewed toward the present and not simply a chance occurrence. Finally, we review reports of fossils that have been assigned to Acanthaceae that are substantially older than the lower Cretaceous estimate for Angiosperms as a whole (i.e., the general consensus that has resulted from several recent dating and fossil-based studies in plants). This is the first study to reconstruct divergence times among clades of Acanthaceae and sets the stage for comparative evolutionary research in this and related families that have until now been thought to have extremely poor fossil resources.

Phylogenetic relationships of Passiflora subgenus Decaloba were examined using 148 taxa and four molecular markers: nuclear nrITS, ncpGS, cp trnL-F, and ndhF. Relationships of subgenus Decaloba to the other four Passiflora subgenera (Astrophea, Deidamioides, Tetrapathea, and Passiflora) are investigated, as are relationships among the eight supersections within subgenus Decaloba. Results indicate that subgenus Deidamioides is not monophyletic. Subgenus Astrophea + subgenus Deidamioides (section Tryphostemmatoides) together form the most basally branching lineage in the genus, followed by a clade comprised of subgenus Passiflora + subgenus Deidamioides (sections Tetrastylis, Polyanthea, and Deidamioides). Passiflora obovata (subgenus Deidamioides section Mayapathanthus) is resolved as part of subgenus Decaloba. The Old World subgenus Tetrapathea is supported as sister to subgenus Decaloba. Subgenus Decaloba is monophyletic and contains seven major lineages that generally correspond to currently re...

Neotropical ecosystems house levels of species diversity that are unmatched by any other region on Earth. One hypothesis to explain this celebrated diversity invokes a model of biotic interactions in which interspecific interactions drive diversification of two (or more) lineages. When the impact of the interaction on diversification is reciprocal, diversification of the lineages should be contemporaneous. Although past studies have provided evidence needed to test alternative models of diversification such as those involving abiotic factors (e.g., Andean uplift, shifting climatological regimes), tests of the biotic model have been stymied by lack of evolutionary time scale for symbiotic partners. In this study, we infer timescales for diversification of hummingbirds and a species-rich plant lineage that is ,50% hummingbird pollinated, Ruellia (Acanthaceae). Results demonstrate that hummingbirds originated about 20 million years before New World Ruellia and that all but one major hummingbird clade was extant before the plant group originated. Thus, the classic model of ‘‘diffuse co-evolution’’ between hummingbirds and this group of plants is rejected by our data. However, together with the observation that the Neotropical clade of Ruellia (,350 species) is far more species rich than its Old World sister group (,75 species), our results are consistent with the hypothesis that plant diversification in the Neotropics has been facilitated in part by a pre-existing diversity of hummingbirds. This hypothesis may find support in other lineages of Neotropical plants that similarly exhibit asymmetrical partitioning of species diversity in the Paleovs. Neotropics.

Phylogenetic knowledge of the large plant family Acanthaceae has been greatly advanced over the last 2 decades. Studies have demonstrated the existence of several major lineages, most of which have been the focus of subsequent investigation. Missing among these is comprehensive study of the 48 genera currently classified in tribe Ruellieae, a pantropical lineage that includes several species-rich genera. We compared the number of validly published names to current estimates of species richness per genus in Ruellieae and found more than 2600 names available for ∼1200 species. Using molecular data from two nuclear (ITS+5.8S, Eif3E) and three chloroplast (trnG-trnR, trnG-trnS, psbA-trnH) markers, we test the placement of these 48 genera in Ruellieae, explore the monophyly of currently recognized taxa, and propose morphological features to diagnose major clades within the tribe. We were able to sample all but four of 48 genera, and all were resolved in Ruellieae except Zygoruellia. Many monospecific or oligos...

As part of our study of the Acanthaceae of Costa Rica and from a phylogenetic study of Ruellia as a whole, we place R. barbillana and R. metallica in synonymy with R. terminalis. We discuss species limits of R. terminalis and provide a key that distinguishes it from close relatives including R. grantii, R. oaxacana, and R. phyllocalyx. Ruellia puri is put into synonymy with R. jussieuoides, and R. standleyi is synonymized with R. ochroleuca. Disjunct distributions characterize R. terminalis, R. jussieuoides, and R. ochroleuca, although this pattern is most extreme in R. ochroleuca. Phylogenetic analysis of DNA sequence data lends support to our synonymy decisions. New lectotypifications are provided for Arrhostoxylum achimeniflorum, Dipteracanthus puri, Dipteracanthus puri β angustifolius, Dipteracanthus puri γ gymnocladus, Lychniothyrsus ochroleucus, and Otacanthus pearcei.

We took a two-tiered approach to test monophyly of Nelsonioideae and place the group within Lamiales, and to determine relationships among taxa within the group. Phylogenetic analysis of a molecula ...

In the early 1990s, a comprehensive set of missions and goals for the discipline was articulated by a global community of systematists; these were presented as Systematics Agenda 2000 (1994). Abbreviated here as SA2K, this agenda spurred awareness of the field and initiated discussions about the role of systematics within biology (e.g., Blackmore and Cutler 1996; Cracraft 2002; Halanych and Goertzen 2009), in education (e.g., Krishtalka and Humphrey 2000; Thanukos 2010), and public policy (e.g., Prance 1995). After nearly 20 years of achievement and growth in systematic biology, a series of four US National Science Foundation-sponsored workshops on “Future Directions in Biodiversity and Systematics Research” was held during 2009–2010 to evaluate progress in the field and identify new directions and opportunities. Workshop participants reviewed SA2K as a way to rapidly achieve common ground and to jump-start our discussions. We did not plan to undertake a formal revision of SA2K, but our discussions led to consensus on a number of relevant points. We share these here with the intention of generating further reflection and discussion toward advancing our field and its missions.

The pace of species discovery and documentation remains too slow on a human-altered planet in the midst of a massive extinction event. Increasing this pace requires altering conventional workflows. In this review, we propose that systematics needs to shift to a model of quantum contributions whereby species hypotheses are published as they are formulated and data as they are collected in web-based repositories and content-management systems. If our recommendation is followed, many species will make their first appearance on the Internet as candidate new species before documentation is complete. Acknowledging the changes that we describe may be controversial, we discuss problems that may be encountered along with possible solutions.

Nichol's turk's head cactus Echinocactus horizonthalonius var. nicholii (Cactaceae) occurs in a few isolated populations in the Sonoran Desert of south-central Arizona (Pima and Pinal counties) and in one isolated population in Mexico. Populations of this variety are disjunct from the more widespread typical variety (E. h. var. horizonthalonius) that occurs in the Chihuahuan Desert of New Mexico, Texas, and Mexico. Variety nicholii occurs almost exclusively on Horquilla limestone substrate, and was federally listed as endangered in 1979. Since 1995, we have monitored study plots of this cactus in the Waterman Mountains, Pima County, Arizona, 32°20′N, 111°28′W, elevation 975 m, measuring size of plants, condition, and presence-absence of reproduction in ca. 200 individuals. Here we report data for 1995–2008. Rates of growth were slow, averaging 0.35 cm (SD ± 0.53) in height and 0.26 cm (SD ± 0.28) in diameter/year. Plants usually began flowering when they reached about 4 cm in height and 8 cm in d...

Stimulated in large part by the advent of the Internet, research productivity in many academic disciplines has changed dramatically over the last two decades. However, the assessment system that governs professional success has not kept pace, creating a mismatch between modes of scholarly productivity and academic assessment criteria. In this article, we describe the problem and present ideas for solutions. We argue that adjusting assessment criteria to correspond to modern scholarly productivity is essential for the success of individual scientists and of our discipline as a whole. The authors and endorsers of this article commit to a number of actions that constitute steps toward ensuring that all forms of scholarly productivity are credited. The emphasis here is on systematic biology, but we are not alone in experiencing this mismatch between productivity and assessment. An additional goal in this article is to begin a conversation about the problem with colleagues in other subdisciplines of biology.

Hybridization in the wild between closely related species is not unusual. In some cases, hybridization may prove beneficial for a rare taxon. Under certain conditions, however, a rare taxon can be driven rapidly to extinction by hybridizing with a more common taxon. This problem is urgent because human activities are increasingly bringing together cross-compatible species that were previously geographically isolated. US conservation policy has yet to address how to deal with hybrid-derived individuals whose ancestry includes an endangered species. Developing sound science-based conservation policy that addresses hybridization requires cross-disciplinary social-science and life-science research to address the following two questions: (1) How do human decisions with regard to species protection, trade, transportation, land use, and other factors affect the opportunities for, and rates of hybridization between, rare species and more common relatives? and (2) How do the positive or negative perceived values regarding hybrids and hybrid-derived individuals compare with values regarding their nonhybridized counterparts from social, cultural, economic, and environmental perspectives? In this article we explore the ways to inform such policy using a multidisciplinary approach.

Acanthaceae as circumscribed today consists of the three subfamilies Acanthoideae (Acanthaceae sensu stricto), Thunbergioideae and Nelsoniodieae, plus the genus Avicennia. Due to the morphological dissimilarities of Thunbergioideae and Nelsonioideae, the delimitation of the family has been controversial. The mangrove genus Avicennia was only recently associated with Acanthaceae for the first time, based on molecular evidence, but without morphological support. In this thesis, phylogenetic analyses of nuclear and chloroplast DNA sequences were used to test the monophyly and exact positions of Thunbergioideae and Nelsonioideae, and to infer detailed phylogenetic relationships within these subfamilies and among major lineages of Acanthaceae. Floral structure and development were comparatively studied in Avicennia and other Acanthaceae using scanning electron microscopy and stereo microscopy. Phylogenetic analyses strongly support monophyly of Thunbergioideae and Nelsonioideae, and place the latter clade with strong support as sister to all other plants treated as Acanthaceae. Thunbergioideae and Avicennia are moderately supported as sister taxa, and together they are sister to Acanthoideae. The general morphology of Avicennia can be easily accommodated in the Acanthaceae, and three synapomorphies support the suggested sister group relationship of Avicennia and Thunbergioideae: (1) collateral ovule arrangement, (2) vertical orientation of ovule curvature, and (3) an exposed nucellus that is contiguous with the ovary wall. Within Thunbergioideae and Nelsonioideae, support values for major lineages are generally high. With some exceptions, the constituent genera are supported as monophyletic. Evolutionary relationships among and within genera are discussed in a morphological and biogeographical context.

We used DNA sequence data from five genic regions (nrlTS; chloroplast trnL-F, trnT–L, rps16, trnS–G) to study phylogenetic relationships of the Tetramerium lineage (Acanthaceae: Justicieae). From a sample of 70 species (representing 25 genera) previously affiliated with the Tetramerium lineage, 68 are included therein. Our analyses excluded Papuasian Calycacanthus and Neotropical Streblacanthus monospermus from the Tetramerium lineage; however, two species described in Justicia (J. gonzalezii and J. medranoi) and a Malagasy species of uncertain generic affinities are nested within the lineage. A monophyletic Tetramerium lineage consists of 23 currently recognized genera with at least 168 species, more than 70% of which occur in the New World. Old World Chlamydocardia and Clinacanthus are serially sister to all other members of the lineage. Other Old World taxa consist of: Ecbolium clade (all sampled species of Ecbolium plus Malagasy Papulina richardii), Megalochlamys clade (Megalochlamys, Trichau...

Phylogenetic relationships of Isoglossinae (Acanthaceae: Justicieae) were studied with a taxon sample including all but one genus putatively placed in the lineage and a dataset composed of DNA sequences from three genic regions (nrITS, cp trnS/G, trnT/L). Two members each of the lineages of Justicieae previously shown to be phylogenetically adjacent to Isoglossinae were included to test monophyly of Isoglossinae. The Malagasy genera Forcipella and Populina were excluded from Isoglossinae by our analysis. The former was placed outside of Justicieae. The latter was placed with plants representing the Tetramerium lineage of Justicieae. Our results do not provide strong support for placement of Ptyssiglottis with Isoglossinae but also cannot refute this placement. Core Isoglossinae (i.e., members of Old World Conocalyx, Isoglossa, Brachystephanus and New World Kalbreyeriella, Stenostephanus s.1., Razisea) are monophyletic; these plants share "Gurtelpollen", hypothesized to be a morphological synapomorphy for the group. Malagasy Conocalyx is part of the clade that includes all sampled species of lsoglossa and does not seem distinct from the latter genus. Sister to Isoglossa (including Conocalyx) is a lineage including all sampled species of Brachystephanus plus all New World Isoglossinae. Plants belonging to this last clade share monothecous stamens as a morphological synapomorphy. Our data do not support but cannot refute reciprocal monophyly of Brachystephanus and New World Isoglossinae. There is, in fact, very little variation among these taxa such that the problem is lack of resolution rather than support for relationships that conflict with current taxonomy. NW plants in particular are richly diverse in morphology and it is remarkable that this diversification is not reflected in variation in these three DNA regions.

We used DNA sequence data from four regions ([1] nrITS; the chloroplast [2] rps16 intron, [3] trnG-S spacer, and [4] trnL-F intron and spacer) to study phylogenetic relationships within tribe Acantheae (Acanthaceae). Our sample includes 18 of 20 recognized genera and 82 of ca. 500 species (plus two Justicieae as out-groups). Results of parsimony and Bayesian analyses were entirely congruent and provided strong support for monophyly of two sub-lineages of Acantheae, referred to here as the ‘one-lipped corolla’ and ‘two-lipped corolla’ lineages, reflecting notable differences in corolla morphology. Subsequent analyses were of the two sublineages separately in order to include all characters (a hypervariable region of trnG-S could not be aligned across the full range of taxa but could be aligned within sublineages). The ‘one-lipped corolla lineage’ comprises six clades of Old World taxa related as follows: [Crossandra (Sclerochiton clade {Cynarospermum [Blepharis (Acanthus clade + Acanthopsis)]})]. ...

We studied nectar characteristics in 12 species of Neotropical hummingbird-pollinated plants: Aphelandra golfodulcensis, A. sinclairiana, A. storkii, Justicia aurea (Acanthaceae); Heliconia irrasa, H. latispatha, H. mathiasii (Heliconiaceae); Pavonia dasypetala (Malvaceae); Isertia haenkeana, Palicourea guianense, Pentagonia macrophylla (Rubiaceae); and Renealmia cernua (Zingiberaceae). The diurnal patterns in sugar content and cumulative volume of nectar were documented in flowers that were protected from flower visitors. Sugar content in these species (mean of all flowers sampled per species ranged from 16 to 28%) was comparable to those reported for other hummingbird-pollinated species. Mean nectar volume per flower spanned an order of magnitude (i.e., 8.8 [Palicourea]–72.7 μl [H. latispatha]), indicating that hummingbirds visited flowers with a wide range of reward. We observed low intraspecific variation in sugar content but high variation in nectar volume. Among species, 10–50 percent of th...

We examined relationships between fragrance and phylogeny using a number of approaches to coding fragrance data and comparing the hierarchical information in fragrance data with the phylogenetic signal in a DNA sequence data set. We first used distance analyses to determine which coding method(s) best distinguishes species while grouping conspecifics. Results suggest that interspecific differences in fragrance composition were maximized by coding as presence/absence of fragrance compounds and biosynthetic pathways rather than when quantitative information was also included. Useful systematic information came from both compounds and pathways and from fragrance emitted by both floral and vegetative tissues. The coding methods that emerged from the distance analyses as best distinguishing species were then adapted for use in phylogenetic analysis. Although hierarchical signal among fragrance data sets was congruent, this signal was highly incongruent with the phylogenetic signal in the DNA sequence data. Notably, topologies inferred from fragrance data sets were congruent with the DNA topology only in the most distal portions (e.g., sister group pairs or closely related species that had similar fragrance profiles were often recovered by analyses of fragrance). Examination of consistency and retention indices for individual fragrance compounds and pathways as optimized onto one of the most-parsimonious trees inferred from DNA data revealed that although most compounds were homoplastic, some compounds were perfectly congruent with the DNA phylogeny. In particular, compounds and pathways found in a few taxa were less homoplastic than those found in many taxa. Pathways that synthesize few volatiles also seem to have lower homoplasy than those that produce many. Although fragrance data as a whole may not be useful in phylogeny reconstruction, these data can provide additional support for clades reconstructed with other types of characters. Factors other than phylogeny, including pollinator interactions, also likely influence fragrance composition. (Acleisanthes; character coding; four o'clocks; fragrance; Mirabilis; phylogenetics; scent; volatiles.)

GC-MS analyses of nocturnal and diurnal floral volatiles from nine tobacco species (Nicotiana; Solanaceae) resulted in the identification of 125 volatiles, including mono- and sesquiterpenoids, benzenoid and aliphatic alcohols, aldehydes and esters. Fragrance chemistry was species-specific during nocturnal emissions, whereas odors emitted diurnally were less distinct. All species emitted greater amounts of fragrance at night, regardless of pollinator affinity. However, these species differed markedly in odor complexity and emission rates, even among close relatives. Species-specific differences in emission rates per flower and per unit fresh or dry flower mass were significantly correlated; fragrance differences between species were not greatly affected by different forms of standardization. Flowers of hawkmoth-pollinated species emitted nitrogenous aldoximes and benzenoid esters on nocturnal rhythms. Four Nicotiana species in section Alatae sensu strictu have flowers that emit large amounts of 1,8 cineole, with smaller amounts of monoterpene hydrocarbons and alpha-terpineol on a nocturnal rhythm. This pattern suggests the activity of a single biosynthetic enzyme (1,8 cineole synthase) with major and minor products; however, several terpene synthase enzymes could contribute to total monoterpene emissions. Our analyses, combined with other studies of tobacco volatiles, suggest that phenotypic fragrance variation in Nicotiana is shaped by pollinator- and herbivore-mediated selection, biosynthetic pathway dynamics and shared evolutionary history.

Heterostyly (i.e., reciprocal placement of anthers and stigmas between two or three floral morphs) is hypothesized to enhance outcrossing and reduce selfing. However, few studies have documented reciprocity among individual plants; instead, mean anther and stigma heights for floral morphs are usually reported, masking interindividual variation. We measured eight floral dimensions for individuals in five populations of three heterostylous Rubiaceae. The three methods used to quantify reciprocity yielded different conclusions regarding the degree to which populations conformed to expectations for heterostylous plants. Only Psychotria poeppigiana had stigma and, to a lesser degree, anther heights in discrete classes. Variation among plants of Bouvardia ternifolia and Psychotria chiapensis yielded a continuum of anther and stigma heights across populations. Comparison of distances between stigma and anthers indicated that only flowers of B. ternifolia had, as expected, a constant value for this distance. Finally, regression relationships between anther and stigma heights and corolla length showed that only in one population each of B. ternifolia and P. poeppigiana, and in P. chiapensis, was distance between anthers and stigmas the same across the range of corolla sizes for both floral morphs. Variation among these species in expression of heterostyly was not clearly linked to phylogenetic relationship or pollinator syndromes. Two approach herkogamous (AH) species were studied for comparison. Flowers of Psychotria brachiata were consistently AH, but flowers of P. pittieri were highly variable. Determining fitness consequences of population-level variation in sexual systems requires studies linking floral morphology to pollinator behavior and pollen transfer.

We used sequence data from the intron and spacer of the trnL-trnF chloroplast region to study phylogenetic relationships among Acanthaceae. This region is more variable than other chloroplast loci that have been sequenced for members of Acanthaceae (rbcL and ndhF), is more prone to length mutations, and is less homoplasious than these genes. Our results indicate that this region is likely to be useful in addressing phylogenetic questions among but not within genera in these and related plants. In terms of phylogenetic relationships, Elytraria (representing Nelsonioideae) is more distantly related to Acanthaceae sensu stricto (s.s.) than Thunbergia and Mendoncia. These last two genera are strongly supported as sister taxa. Molecular evidence does not support monophyly of Acanthaceae s.s., although there is strong morphological evidence for this relationship. There is strong support for monophyly of four major lineages within Acanthaceae s.s.: the Acanthus, Barleria, Ruellia, and Justicia lineages as here defined. The last three of these comprise a strongly supported monophyletic group, and there is weaker evidence linking the Ruellia and Justicia lineages as closest relatives. Within the Acanthus lineage, our results confirm the existence of monophyletic lineages representing Aphelandreae and Acantheae. Lastly, within the Justicia lineage, we develop initial hypotheses regarding the definition of sublineages; some of these correspond to earlier ideas, whereas others do not. All of these hypotheses need to be tested against more data. Acanthaceae are a large family (.4000 species; Mabberley, 1987), the members of which present a rich diversity of morphological and ecological characteristics. The family is part of Lamiales s.l. (sensu lato) (i.e., sensu Olmstead et al., 1993). Unlike a number of family-level taxa in Lamiales (e.g., Lamiaceae, Verbenaceae, Scrophulariaceae), there seems little doubt that Acanthaceae s.s. (sensu stricto, see below) are monophyletic. This hypothesis is supported by the shared presence of a fruit type that is unique among angiosperms: a few-seeded, explosively dehiscent capsule within which the seeds are borne on retinacula (the lignified derivatives of funiculae). The precise delimitation of the family, however, has been controversial due to three small lineages that do not 1

A survey of bracteal (extrafloral) nectaries in species of Aphelandra (Acanthaceae) reveals substantial diversity. Each bracteal nectary is an aggregate of individual glands that vary in number, size, and structure among species. Glands contain three cell layers: a palisade-like secretory cell layer, a one-to-many-celled intermediate layer with thickened cell walls, and a foot layer. Members of the A. pulcherrima complex have one of two distinct gland types: relatively small glands with a single-celled intermediate layer or larger glands that have a multicellular intermediate layer. Nectaries composed of small glands are patches of many (>50) glands, whereas those composed of large glands are patches of

Much has been made of the current inability to increase the pace of discovery and documentation of the rich pageant of life on this planet [1]. One of the most contentious issues is whether new publishing practices, especially utilization of web-based knowledge repositories and publishing platforms, are panacea, placebo, or anathema ([2,3] in support; [4,5] against). In a recent TREE article [6], we posited that the pace of discovery and documentation could be increased by changing the current practice of publishing data and species hypotheses.

Acanthaceae (Asteridae; Lamiales) include ∼4000 species and encompass a range of morphological diversity, habitats, and biogeographic patterns. Although they are important components of tropical and subtropical habitats worldwide, inadequate knowledge of the family's phylogenetic framework has impeded comparative research. In this study, we sampled all known lineages of Acanthaceae including Andrographideae. Also included were eight of 13 genera whose relationships remain enigmatic. We used sequence data from nrITS and four chloroplast noncoding regions, and parsimony and Bayesian methods of analysis. Results strongly support most aspects of relationships including inclusion of Avicennia in Acanthaceae. Excepting Neuracanthus, newly sampled taxa are placed with strong support; Kudoacanthus is in Justicieae, Tetramerium lineage, and the remaining enigmatic genera are in Whitfieldieae or Barlerieae, and Andrographideae are sister to Barlerieae. This last result is unanticipated, but placement of Andrographideae based on structural characters has been elusive. Neuracanthus is monophyletic but placement relative to (Whitfieldieae (Andrographideae + Barlerieae)) is weakly supported. Many clades have clear morphological synapomorphies, but nonmolecular evidence for some remains elusive. Results suggest an Old World origin with multiple dispersal events to the New World. This study informs future work by clarifying sampling strategy and identifying aspects of relationships that require further study.

ummingbirds and the flowers they visit are among the most colorful examples of mutualistic relationships between animals and flowering plants. The acrobatics, pugnacious behavior, and startling colors of many hummingbird species have won them the attention of biologists and hobbyists alike. In turn, the large and showy flowers of many of the plants pollinated by hummingbirds make them favorites of horticulturalists and well-collected by botanists. I have studied the systematics and reproductive biology of the neotropical plant genus Aphelandra (Acanthaceae). These plants have long, tubular, brightly colored (yellow, orange, red, and pink), and nectar-rich corollas that are the epitome of the hummingbird-pollinated flower (Faegri and van der Pijl 1966). From more than 100 hours spent observing visitors to Aphelandra flowers, I have documented that they are indeed pollinated by hummingbirds. Most of the plant species have strongly curved corollas that are 5.5-7 cm long (Figure la-d); these are pollinated by hermit hummingbirds (subfamily Phaethorninae), whose long, decurved bills match the

A latitudinal gradient in biodiversity has existed since before the time of the dinosaurs, yet how and why this gradient arose remains unresolved. Here we review two major hypotheses for the origin of the latitudinal diversity gradient. The time and area hypothesis holds that tropical climates are older and historically larger, allowing more opportunity for diversification. This hypothesis is supported by observations that temperate taxa are often younger than, and nested within, tropical taxa, and that diversity is positively correlated with the age and area of geographical regions. The diversification rate hypothesis holds that tropical regions diversify faster due to higher rates of speciation (caused by increased opportunities for the evolution of reproductive isolation, or faster molecular evolution, or the increased importance of biotic interactions), or due to lower extinction rates. There is phylogenetic evidence for higher rates of diversification in tropical clades, and palaeontological data demonstrate higher rates of origination for tropical taxa, but mixed evidence for latitudinal differences in extinction rates. Studies of latitudinal variation in incipient speciation also suggest faster speciation in the tropics. Distinguishing the roles of history, speciation and extinction in the origin of the latitudinal gradient represents a major challenge to future research.

We present results of dynamic head-space collections and GC–MS analyses of floral and vegetative fragrances for 20 species in three genera of Nyctaginaceae: Acleisanthes, Mirabilis and Selinocarpus. Most of the species included in this study are either hawkmoth or noctuid moth-pollinated. A wide variety of compounds were observed, including mono- and sesquiterpenoids, aromatics (both benzenoids and phenylpropanoids), aliphatic compounds, lactones, and nitrogen-bearing compounds. Intraspecific variation in fragrance profiles was significantly lower than interspecific variation. Each species had a unique blend of volatiles, and the fragrance of many species contained species-specific compounds. The fragrance profiles presented here are generally consistent with previous studies of fragrance in a variety of moth-pollinated angiosperms.

I examined three aspects of the cladistic treatment of a set of 17 F1 hybrids of known parental origin: (1) impact of hybrids on consistency index (CI) and number of most parsimonious trees (Trees), (2) placement of hybrids in cladograms, and (3) impact of hybrids on hypotheses of relationship among species. The hybrids were added singly and in randomly selected sets of two to five to a data set composed of Central American species of Aphelandra (including the parents of all hybrids). Compared to analyses with the same number of OTUs all of which were species, the analyses with hybrids yielded results with significantly higher CI. There was no difference in Trees between analyses with hybrids versus species. There was thus no evidence that hybrids would appear to be more problematic for cladistic methods than species. Accordingly, hybrids will not be readily identifiable as taxa that cause marked change in these indices. About % of the hybrids were placed as the cladistically basal members of the lineage that included the most apomorphic parent. Relatively apomorphic hybrids were placed proximate to the most derived parent (ca. 13% of hybrids). Other placements occurred more rarely. The most frequent placements of hybrids thus did not distinguish them from normal intermediate or apomorphic taxa. When analyses with hybrids yielded multiple most parsimonious trees, these were no more different from each other than were the equally parsimonious trees that resulted from analyses with species. Most analyses with one or two hybrids resulted in minor or no change in topology. When hybrids caused topological change, they frequently caused rearrangements of weakly supported portions of the cladogram that did not include their parents. When they disrupted the cladistic placement of their parents, they often caused their parents to change positions, with at least one topology bringing the parental lineages into closer proximity with the hybrid placed between them. Hybrids between parents from the two main lineages of the group caused total cladistic restructuring. In fact, the degree of relationship between a hybrid's parents (measured by both cladistic and patristic distance) was strongly correlated with CI (negatively) and with the degree of disturbance to cladistic relationships (positively). Thus, hybrids between distantly related parents resulted in cladograms with low CI and major topological changes. This study suggests that hybrids are unlikely to cause breakdown of cladistic structure unless they are between distantly related parents. However, these results also indicate that cladistics may not be specially useful in distinguishing hybrids from normal taxa. The applicability of these results to other kinds of hybrids is examined and the likely cladistic treatment of hybrids using other sources of data is discussed.