Contrasting patterns of postglacial range shifts between the northern and southern hemisphere in Herbertus (Herbertaceae,Marchantiophyta)

The leafy liverwort genus Herbertus exhibits considerably variable morphology and widely disjunct distributions in both hemispheres. Here, the biogeographic history of the genus and its phylogenetic relationships with the focus on the taxonomically difficult, northern hemispheric disjunct species, were investigated. We conducted a time-calibrated, molecular-based phylogenetic analysis using psbA, trnL-F, and ITS1-2 loci and different approaches for ancestral range inference of the genus. Herbertus is inferred to have originated in the Cenozoic era about 51 million years ago, in an ancestral area including southern South America, the Neotropics, Oceania, and South-east Asia. The current distribution of the genus is supported to have resulted from long-term in situ persistence, short and long distance dispersals, extinctions and recolonizations. Postglacial range shifts between the southern and northern hemisphere in the genus show distinct patterns. In the southern hemisphere, Herbertus is characterized by in situ persistence, not showing further dispersal until the uplift of the Andean Cordillera. Species of the northern hemisphere showed wide range expansions and repeated recolonizations, including north- and southward dispersals, recolonizations and extinctions. Our results support that the ancestor of South-east Asiatic Herbertus had a Gondwanan origin and arrived in Asia via Indian Plate migration. The uplift of high mountains must have had a strong influence in the diversification and dispersal of the genus. Our results further suggest that climate changes must have had a profound effect on the evolution and biogeography of the species of Herbertus in the northern hemisphere, and might also have influenced the reproductive strategies of the genus. Few genetic differentiations amongst currently recognized species H. aduncus, H. dicranus, H. hutchinsiae, H. stramineus, H. delavayi, and H. kurzii, and amongst H. sendtneri, H. armitanus, and H. circinatus were shown, suggesting that the morphological characters that are currently used for delimiting species should be re-evaluated.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF SOUTHEASTERN NORTH AMERICA: PHILODENDRON LEAF REMAINS

Large leaves, new to the fossil record, from the Claiborne Formation in western Tennessee have been collected and analyzed. Careful analyses of venation and cuticular anatomy indicate that these fossil leaves contain specific characters found in modern species of the genus Philodendron subgenus Meconostigma. Features of venation were taxonomically more useful than other features. Reports of fossils of the Araceae are scarce and those few fossils assigned to the family frequently have been inadequately studied. This report establishes a reliable and carefully documented occurrence of the Araceae in the fossil record, and provides information concerning the differentiation of the family in time and the distributions of the genus Philodendron.     

A taxonomic revision and pollen morphology of the genus Dendrokingstonia (Annonaceae)

The genus Dendrokingstonia (Annonaceae) is taxonomically revised and palynologically studied. Three species are recognized, one of which, D. gardneri , is described as new to science. One new combination, D. acuminata , is made. The genus occurs from southern Thailand to Peninsular Malaysia and Sumatra. On the basis of macromorphology and pollen characters, it is considered to be related to Monocarpia. Both genera show a combination of macromorphological characters that is rare in the family, i.e. considerably enlarged stigmas, leaves with percurrent tertiary veins, a highly reduced number of carpels per flower and relatively large monocarps with a thick, hard wall. Scanning and transmission electron microscopy show that the pollen grains of Dendrokingstonia and Monocarpia are monosulcate monads with a columellate infratectum and a more or less bulging intine at the sulcus. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 76–90.     

Pollen morphology of Strobilanthes Blume (Acanthaceae) in China and its taxonomic implications

The pollen morphology of 37 Chinese species of Strobilanthes Blume (Acanthaceae), a large and taxonomically problematic genus, was investigated by LM and SEM. Two distinct pollen shape classes, ellipsoid and spheroidal, recognised by earlier workers were found. Eight pollen types could be distinguished on the basis of variation in ornamentation, and the number, distribution and type of apertures. Five of these fall within twenty two pollen types reported from Strobilanthes in southern India and Sri Lanka. Three new pollen types are also described, further extending the palynological diversity of the genus. From the results presented, the characters of palynological diversity and the related taxonomic implications are discussed.     

Revision of the genus Rhotala Walker, 1857 (Hemiptera: Fulgoroidea: Achilidae) in China,with description of a new species

The achilid genus Rhotala is taxonomically revised based on specimens from China. Four species are recognized in this paper as valid: R. vittata Matsumura, 1914; R. dimidiata Jacobi, 1944; R. fanjingshana Chen et Yang, 2008; and one new species: R. jiangxiensis sp. nov. (southern China: Jiangxi). Male genitalia of these species are described and illustrated. In addition, hind tibia, forewings and hindwings are illustrated for the male of R. jiangxiensis sp. nov. A diagnosis of the genus is redefined. A key to the species of Rhotala in China is presented.     

Consistent and variable leaf anatomical characters in Carex (Cyperaceae)

The anatomy and morphology of leaves in Carex have the potential to be taxonomically useful. However, studies on the variability of leaf characteristics in the genus are sparse. Researchers therefore risk using leaf anatomical characters without the knowledge of whether they are consistent in a species. We examined 22 qualitative and seven quantitative leaf anatomy characters from transverse leaf sections to test their consistency across 11 Carex spp. The characters were clearly described and primarily microscopic. Some characters were found to exhibit high levels of intraspecific variation, whereas other characters exhibited high levels of consistency in a species, including the shape of the leaf section, the density of papillae and the size of epidermal cells. Caution must be applied when choosing leaf anatomy to delimit taxa because of the intraspecific variability found in some characters, but sufficient numbers of invariant characters exist to provide useful taxonomic separation. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 371–384.     

Morphological studies on the nutlet of Turkish Cyclotrichium Manden. & Scheng. (Lamiaceae)

Morphological characters of the nutlets of Cyclotrichium Manden. & Scheng. in Turkey were examined using scanning electron microscopy (SEM). The genus Cyclotrichium is represented by six species in Turkey, of which two are endemic for Turkey. Nutlet morphology of the species was studied using stereoscopic microscopy and SEM, and the features were described and illustrated. Nutlet morphology of the examined specimens exhibits some variation in size, shape and sculpture. Nutlets range from 1.1 to 2.0 mm in length and 0.5 to 1.0 mm in width and are elliptic, oblong to ovoid and trigonous in shape. Nutlet sculpture can be divided into two main types: distinctly or weakly reticulate pattern in C. niveum, C. origanifolium, C. longiflorum and rounded cell arrangement in the remaining species. Micromorphological characters could be useful in solving taxonomic problems of Cyclotrichium.     

Revision of the Neotropical Genus Hemibidessus Zimmermann (Coleoptera: Dytiscidae: Hydroporinae: Bidessini)

The genus Hemibidessus Zimmermann, 1921 is revised. Six species are recognized in the genus. Two new species are described, H. spirodiscus sp. n. from Bolivia and H. spangleri sp. n. from Argentina. A key for identifying the species is presented. The female genitalia are thoroughly illustrated and described for the first time for any species of Bidessini. Other important diagnostic structures are also illustrated including the male genitalia. A cladistic analysis is presented for 8 species (6 ingroup and 2 outgroup species) and 13 characters. A single most parsimonious cladogram was found.     

The ontogeny of appendages of Heterocypris salina (Brady, 1868) Ostracoda (Crustacea)

The post-embryonic development of the appendages of the Cyprididae ostracod Heterocypris salina (Brady, 1868) are described in detail and compared with those of other podocope species documented in previous studies. Generally, the appearence of limbs during onotgeny of H. salina is similar to that of other species, but small differences in limb morphologies were identified between H. salina and other Cyprididae species, including other Heterocypris species. Some features appear either earlier or later in the development of H. salina compared with other species, even species of the same genus. These features may be useful characters for phylogenetic analyses at the genus and family levels.     

Nutlet micromorphology of Iranian Nepeta (Lamiaceae) species

To evaluate nutlets characteristics for systematic relationships, a comprehensive morphological and micro‐morphological study of the nutlets of 16 Iranian taxa of Nepeta (Lamiaceae) was conducted using scanning electron microscopy (SEM). Differences in surface ornamentation, size, shape and color were observed between the species. The studied taxa were categorized in two basic types based on surface ornamentation: non‐sculptured (smooth‐type) and sculptured nutlets. The non‐sculptured nutlets type could be further divided into four sub‐types, including smooth, granulate, undulate‐cellular and reticulate. In the sculptured nutlets, three subtypes of tuberculate, tuberculate‐cellular and verrucate were recognized. The shape of nutlets were described as oblong, ovoid‐oblong, oblong‐linear, to elliptic‐oblong and their size range are 1.2–2.0 mm in length and 0.5–2.0 mm in width. Based on micrographs, the areole form and location were defined as bi‐lobed, straight, basal, sub‐basal or lateral. Nutlet micromorphological characteristics such as surface ornamentation can be useful for classification and identification of e.g. medicinal species of Nepeta in Iran.     

Nutlet micro‐morphology of the genus Microula (Boraginaceae) from the Qinghai–Tibetan Plateau,and its systematic implications

The micro‐morphology of nutlets in 30 taxa (28 species and 2 varieties) of Microula Benth. (Boraginaceae) was surveyed by light and scanning electron microscopy. A principal coordinate‐based ordination analysis (PCO) (8 morphological and 14 micro‐morphological characters) was applied to explore whether taxa clustered according to the current sectional taxonomy. The nutlet epidermis showed three major features: trichomes, papillae and lamellae. Two basic trichome types could be identified: branched and unbranched. Stellate trichomes (STT) and stipitate stellate trichomes (SST) can be considered as two subtypes of branched trichomes based on shape and branching location. Three kinds of unbranched trichomes were recognized by their shapes: slender trichomes (SLT), spear‐like trichomes (SPT) and conical trichomes (CT). Four types of papillae were revealed in this genus: central apiculate papillae (CAP), debris‐covered convex papillae (DCP), rugous papillae (RP) and tabular papillae (TP). The lamellae was classified in five types: glossy lamellae (GL), thread‐like lamellae (TL), rugous lamellae (RL), interrupted lamellae (IL) and debris‐covered lamellae (DCL). The variation in the nature of surface sculpturing, nutlet shape and size, dorsal aperture shape, and attachment scar positions proved useful diagnostic characters. Taxa belonging to the same section generally clustered together in the PCO analysis. A key based on the morphology and micro‐morphology of nutlets is provided to distinguish the sections of Microula. The presence of SLT, STT, DCP and DCL is consistent with the macro‐morphological classification and provides additional evidence for the subgeneric delimitations. The SST, SLT, CAP, DCP, RP, DCL, GL and RL are useful in discriminating species in Microula. Additionally, the distribution of some trichomes, papillae and lamellae types is correlated with the altitudinal distribution of the species.     

EVALUATION OF RELATIONSHIPS WITHIN ERIODICTYON (HYDROPHYLLACEAE) USING TRICHOME CHARACTERISTICS

All 13 taxa of Eriodictyon Bentham (Hydrophyllaceae) were examined to determine the range of variation in trichome characters within the genus. Four simple trichome types were found: short and straight, intermediate length and straight, long and straight, long and wavy. Glandular capitate trichomes were also found in some species. Sessile glands were also observed but not included in the study. Most taxa displayed unique combinations of trichome types on stems, leaves, inflorescence axes and flower parts that allowed those taxa to be identified using trichome types alone. Trichome data support most previous taxonomic treatments, but suggest that 1) the two varieties of E. traskiae are extremely similar, 2) the rare E. altissimum is most similar to the widespread E. californicum, 3) the rare E. capitatum is allied with E. crassifolium var. nigrescens or perhaps E. angustifolium, 4) E. crassifolium var. denudatum and E. crassifolium var. nigrescens are essentially indistinguishable using trichome characters and, coupled with variation in other characters, are best combined under the name E. crassifolium var. nigrescens. Trichome characters provide a wealth of taxonomically useful information and may prove useful in the study of related genera of Hydrophyllaceae.     

Pollen morphology of the East Asiatic genus Chelonopsis (Lamioideae: Lamiaceae) and allied genera,with reference to taxonomic implications and potential pollination ecology

Pollen grains of 15 taxa of the genus Chelonopsis (14 spp. and 1 variety) from China and Japan and 6 species of the closely related genera Bostrychanthera (1 species) and Gomphostemma (5 species) were examined by light and scanning electron microscopy. Of these, the pollen morphology of 18 taxa was studied for the first time. Pollen grains were found to be tricolpate with polar lengths of 20.8–30.0 μm and equatorial widths of 17.5–27.3 μm. The basic shape of the pollen in most taxa is subprolate or prolate-spheroidal, but spheroidal, subprolate-spheroidal, oblate-spheroidal, and prolate-subprolate grains are also found in some species. The surface is generally reticulate or suprareticulate in Chelonopsis and granulate in Bostrychanthera. In comparison with those of Chelonopsis and Bostrychanthera, the pollen grain surfaces of Gomphostemma are more diverse. In Chelonopsis, pollen is taxonomically useful at the sectional level, and some grains provide enough characters for species delimitation. The potential pollination ecology of cellular hairs on the anthers of Chelonopsis and Bostrychanthera is also briefly discussed.     

Exploring the Afromontane centre of endemism: Kniphofia Moench (Asphodelaceae) as a floristic indicator

Aim The genus Kniphofia contains 71 species with an African–Malagasy distribution, including one species from Yemen. The genus has a general Afromontane distribution. Here we explore whether Kniphofia is a floristic indicator of the Afromontane centre of endemism and diversity. The South Africa Centre of diversity and endemism was explored in greater detail to understand biogeographical patterns. Location Africa, Afromontane Region, southern Africa, Madagascar and Yemen. Methods Diversity and endemism for the genus were examined at the continental scale using a chorological approach. Biogeographical patterns and endemism in the South Africa Centre were examined in greater detail using chorology, phenetics, parsimony analysis of endemicity (PAE) and mapping of range‐restricted taxa. Results Six centres of diversity were recovered, five of which are also centres of endemism. Eight subcentres of diversity are proposed, of which only two are considered subcentres of endemism. The South Africa Centre is the most species‐rich region and the largest centre of endemism for Kniphofia. The phenetic analysis of the South Africa Centre at the full degree square scale recovered three biogeographical areas that correspond with the subcentres obtained from the chorological analysis. The PAE (at the full degree square scale) and the mapping of range‐restricted taxa recovered two and six areas of endemism (AOEs), respectively. These latter two approaches produced results of limited value, possibly as a result of inadequate collecting of Kniphofia species. Only two AOEs were identified by PAE and these are embedded within two of the six AOEs recovered by the mapping of range‐restricted taxa. All the above AOEs are within the three subcentres found by chorological and phenetic analysis (at the full degree square scale) for the South Africa Centre. Main conclusions The centres for Kniphofia broadly correspond to the Afromontane regional mountain systems, but with some notable differences. We regard Kniphofia as a floristic indicator of the Afromontane Region sensu lato. In southern Africa, the phenetic approach at the full‐degree scale retrieved areas that correlate well with those obtained by the chorological approach.     

VARIATION IN NUTLET MORPHOLOGY OF LIMNANTHES

Morphological features of nutlets of all members of the genus Limnanthes were examined by light and scanning electron microscopy (SEM). Eight distinct nutlet types are recognized on the basis of presence or absence, size, shape, density, and color of tubercles and associated filamentous or globular background structures. Some taxa (e.g., L. macounii, L. bakeri) display only one nutlet type, some (e.g., L. floccosa floccosa) have different types in different populations, and others (e.g., L. alba, L. striata) have two or more types within populations. Four classes of nutlets are designated as subtypes on the basis of their forms varying between the eight types recognized. The relationships among various taxa on the basis of nutlet types are schematically diagrammed and compared with other classification schemes. Results suggest that the proposed details of nutlet morphology should be considered in an identification key. Preliminary evidence indicates that certain highly sculptured (tuberculate) nutlet types with large surface areas occur more often in wetter habitats and that variation in this trait in different Limnanthes species might be adaptively associated with geographical distribution patterns.     

Phylogeny of the tribe Abrotrichini (Cricetidae,Sigmodontinae): integrating morphological and molecular evidence into a new classification

The tribe Abrotrichini (five genera and 14 living species) is a small clade within the speciose subfamily Sigmodontinae (Rodentia, Cricetidae), representing one of the extant successful radiations of mammals at southern high latitudes of the Neotropics. Its distribution is mostly Andean, reaching its greatest diversity in southern Argentina and Chile. We evaluate the phylogenetic relationships within this tribe through parsimony and Bayesian approaches based on 99 morphological characters (including 19 integumental characters, 38 skull characters, 31 dental characters, three postcranial skeletal characters, seven from the male accessory glands and phallus and one from the digestive system) and six molecular markers (one mitochondrial and five nuclear). We include representatives of all, except one, of the currently recognized species of living Abrotrichini plus one fossil form. Based on total evidence, we recovered a primary division between the genus Abrothrix and a group including the long‐clawed Abrotrichini, Chelemys, Geoxus, Notiomys and Pearsonomys. Both clades are recognized and named here as subtribes. The large degree of morphological variation observed within Abrothrix suggests that species in the genus fall into four groups, which we recognize as subgenera. In addition, the two known species of Chelemys do not form a monophyletic group, and Geoxus was recovered as paraphyletic with respect to Pearsonomys. To reconcile classification and phylogenetics, we describe a new genus for Chelemys macronyx and include Pearsonomys as a junior synonym of Geoxus. Our results highlight the importance of both morphology and molecules in resolving the phylogenetic relationships within this tribe. Based on biogeographical analyses, we hypothesize that Abrotrichini originated in south‐western South America by vicariance and then diversified mostly by successive dispersal events.     

Taxonomic review of the genus Symmorphus Wesmael (Hymenoptera: Vespidae: Eumeninae) from the Far East

Far Eastern species of the genus Symmorphus Wesmael are taxonomically reviewed. Fourteen species are recognized. Taxonomic information regarding this group that has been treated fragmentarily is summarized in this study. A revised key is presented, and recognition characteristics and images are supplied for each species.