High species diversity in one of the dominant groups of spiders in East African montane forests (Araneae: Pholcidae: Buitinga n. gen., Spermophora Hentz)

The six-eyed pholcid spiders of East Africa are revised. A new genus is recognized, Buitinga n. gen ., with 17 new species and three species transferred from Spermophora Hentz. The new genus is characterized by the presence of a scape on the epigynum. This scape may be straight or tightly curled up at rest, and is usually highly expandable. Seven additional African and Comoran species are newly described and tentatively assigned to Spermophora . A data matrix with 60 characters and 77 taxa (including 20 East African species and 25 additional Spermophora and ' Spermophora -like' species) is analysed cladistically. Buitinga is closer to the genera Paramicromerys Millot (endemic in Madagascar) and Spermophorides Wunderlich (Mediterranean and Canary Islands) as well as to several African and Comoran species tentatively assigned to Spermophora than to the type species of Spermophora . It is argued that current estimates of species numbers may be inaccurate and that pholcids may turn out to be one of the most diverse spider families.  © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society, 2003, 137, 555−619     

Phylogeny of pholcid spiders (Araneae: Pholcidae): combined analysis using morphology and molecules

The spider family Pholcidae comprises a large number of mainly tropical, web-weaving spiders, and is among the most diverse and dominant spider groups in the world. The phylogeny of this family has so far been investigated exclusively using morphological data. Here, we present the first molecular data for the family analyzed in a phylogenetic context. Four different gene regions (12S rRNA, 16S rRNA, cytochrome c oxidase subunit I, 28S rRNA) and 45 morphological characters were scored for 31 pholcid and three outgroup taxa. The data were analyzed both for individual genes, combined molecular data, and molecular plus morphological data, using parsimony, maximum likelihood, and Bayesian methods. Some of the phylogenetic hypotheses obtained previously using morphology alone were also supported by our results, like the monophyly of pholcines and of the New World clade. On the other hand, some of the previous hypotheses could be discarded with some confidence (monophyly of holocnemines, the position of Priscula), and still others need further investigation (the position of holocnemines, ninetines, and Metagonia). The data obtained provide an excellent basis for future investigations of phylogenetic patterns both within the family and among spider families.     

Revision and cladistic analysis of the jumping spider genus Onomastus (Araneae: Salticidae)

The jumping spider genus Onomastus Simon, 1900 is revised. Four new species: Onomastus indra sp. nov. , Onomastus kaharian sp. nov. , Onomastus pethiyagodai sp. nov. , and Onomastus rattotensis sp. nov. are described. Parsimony analysis of 26 morphological characters supported the monophyly of Onomastus. Lyssomanes is sister to Onomastus. Onomastus separates into two clades: the widespread South‐East Asia clade and the South Asia clade. The South Asia clade is restricted to the Sri Lanka–Western Ghats biodiversity hotspot. Species of the South Asia clade appear to be spot endemics, highly in danger of extinction because of habitat loss and climate change. Male palps are complex and species‐specific, suggesting rapid divergent evolution. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 711–745.     

Molecular taxonomy in pholcid spiders (Pholcidae, Araneae): evaluation of species identification methods using CO1 and 16S rRNA

The identification of species using molecular characters is a promising approach in alpha taxonomy and in any discipline depending on reliable assignment of specimens. Previous studies have shown the feasibility of the method, but considerable controversy persists. In this study, we use pholcid spiders in an effort to address two main issues. First, we evaluate and calibrate molecular species (re-)identification within a closely related group of organisms by using specimens that are morphologically unambiguously either conspecific or not. Species limits hypothesized a priori based on morphology were almost universally reconstructed by both mitochondrial markers used. Second, we focus on species identification methodology in a morphology-calibrated scenario, i.e. on how to assess the quality of a dataset and of the method used to obtain distance estimates (e.g. choice of markers, alignment strategy, type of distance data). We develop a number of statistical estimators permitting the measurement and communication of the clarity of species boundaries in a dataset and discuss their benefits and drawbacks. We propose that box plots rather than histograms are the superior tool for graphically illustrating taxonomic signal and that the median is a more appropriate measure of central tendency than the mean. Applying the suggested tools to our data, we propose that in molecular species identification, indel-related alignment uncertainties may often be even advantageous (by accentuating taxonomy-relevant information) and we conclude that — at least for our dataset — 16S is better suited to taxonomy than CO1.     

On American 'Micromerys' and Metagonia (Araneae, Pholcidae), with notes on natural history and genital mechanics

Two American pholcids previously assigned to Micromerys Bradley are redescribed and transferred to Mefagonia: Me. conica (Simon, 1893) comb. n. (Venezuela), and Me. delicata (Cambridge, 1895) comb. n. (Central America). Two close relatives of Me. delicafa are newly described: Me. uvita sp. n. and Me. falamanca sp. n. (both from Costa Rica). The name Me. turrialba Gertsch. 1986 is synonymized with Me. rica Gertsch, 1986 (Costa Rica) which is redescribed. Four additional representatives of Metagonia are described as new: Me. reventazona sp. n. (Costa Rica, Panama), Me. toro sp. n. (Panama), Me. hitoy sp. n. and Me. hondura sp. n. (Costa Rica). Leptopholcus dalei (Petrunkevitch, 1929) from Puerto Rico, which was originally described as Micromerys dalei. is redescribed and its closeness to 'true' Old World Lepfopholcus is supported. Arguments are presented to justify the transfer of the American 'Micromerys' species to Metagonia. Notes on the natural history of Me. rica are presented, and a detailed account on copulation and genital mechanics is given as a basis for future comparison. © 1997 The Norwegian Academy of Science and Letters     

On the distinction between Modisimus and Hedypsilus (Araneae, Pholcidae), with notes on behavior and natural history

Evidence is presented that the genera Hedypsilus Simon, 1893 and Modisimus Simon, 1893 are subjective synonyms. Precedence is given to Modisimus over Hedypsilus. The type species of both genera (M. glaucus Simon, 1893 and H. culicinus Simon, 1893) are redescribed. Hedypsilus lawrencei Lessert, 1938 is synonymized with Hedypsilus culicinus Simon, 1893. It appears to be a pantropical, synanthropic species. Preliminary notes on its natural history are presented. Two closely related new species are described, Modisimus simoni sp. n. (Venezuela) and Modisimus david sp. n. (Central America). Modisimus globosus Schmidt, 1956 (Colombia?) is redescribed and illustrated for the first time. Notes on the usefulness of the curvature of eye rows and of leg formulae for the systematics of pholcids are given.     

Functional morphology of female goblin spider genitalia (Arachnida: Araneae: Oonopidae) with notes on fertilization in spiders

The genital structures of most spiders are poorly investigated in respect of their functional morphology because the traditional taxonomic practice is to inspect slide-mounted genitalia only. The present study describes the female genitalia of three members belonging to the megadiverse haplogyne spider family Oonopidae by means of histological serial sections, scanning electron microscopy, and X-ray ultramicroscopy. The female genitalia of Neoxyphinus ogloblini, Dysderina sp., and Heteroonops spinimanus are complex and might have evolved under sexual selection by cryptic female choice. However, there is no direct evidence for cryptic female choice in these species based on the results of the present study. In N. ogloblini and Dysderina sp., spermatozoa and secretion are stored in a large receptaculum. Highly elongated gland cells filled with secretory vesicles extend over the receptaculum of N. ogloblini. In addition, sperm are present in the uterus internus of female N. ogloblini and Dysderina sp. The location of fertilization is still unknown for most spiders. One female of Dysderina sp. had sperm in the uterus and ovary strongly suggesting that fertilization in this species takes place in the ovary. An anterior sclerite with attached muscles should serve females to lock the uterus externus during copulation as suggested for other oonopids. The male palp of N. ogloblini shows a simple embolus whereas the embolus of Dysderina sp. is more complicated and accompanied by a cork-screw-shaped conductor. Females of H. spinimanus have an anterior sclerite in which thread-like gland ducts lead. The chitinized posterior diverticulum shows peculiar papillae in its anterior wall. The exact location of sperm storage in H. spinimanus remains unknown since spermatozoa were not present in the anterior sclerite and the posterior diverticulum. The anterior sclerite might be used to lock the uterus externus similar to N. ogloblini and Dysderina sp. H. spinimanus was previously suggested to be parthenogenetic and a male has only been recently associated with this species. The male was not investigated for this study.     

Pulchellaranea pedunculatan. gen.n. sp. (Araneae: Araneidae), a new genus of spiders with a review of araneid spiders in Cenozoic Dominican amber

Pulchellaranea pedunculatan. gen., n. sp. (Araneae: Araneidae), a new genus of orb web spiders, is described from Cenozoic Dominican amber. Fossil representatives of the Araneidae are uncommon, even though the family is widely distributed today. An arboreal ant adjacent to Pulchellaranea pedunculata indicates the likely habit of the fossil spider in the Dominican amber forest. The encounter with the ant may have resulted in both specimens falling into fresh resin on the trunk of the Algarroba tree. A key to araneid spiders reported in Dominican amber is provided.http://www.zoobank.org/urn:lsid:zoobank.org:pub:D7F92A3C-46BB-48C0-9BA1-0496038F04A5     

Wahlenbergia cordiformis sp. nov. (Campanulaceae: Campanuloideae), a new woody species from Western Cape,South Africa

Wahlenbergia cordiformis, a new species of the Lightfootia‐type is described and illustrated. It is currently only known from the farm Knolfontein, in the Swartruggens region, north east of Ceres in the Western Cape, South Africa. The closest relative of W. cordiformis is unclear, but it may be closely related to either W. tenella, W. nodosa, W. tenerrima, W. oligantha or W. neorigida. Wahlenbergia cordiformis is distinguished from them by its cordate, leaf‐like calyx lobes.     

Morphology and molecular phylogeny of Oxytricha seokmoensis sp. nov. (Hypotrichia: Oxytrichidae), with notes on its morphogenesis

A new hypotrichous ciliate, Oxytricha seokmoensis sp. nov., was discovered in a soil from a forest in South Korea and described based on the observations of living and stained specimens. In addition, phylogenetic analyses were performed using the small subunit ribosomal RNA (18S rRNA) gene sequence. Morphologically, the new species is similar to the O. granulifera-complex in terms of ciliary structure and arrangement of cortical granules, but dorsal kineties 3 and 4 (not completely separated vs. separated) and macronuclear nodules in the cyst (separated vs. fused) differ. Oxytricha seokmoensis is most similar to O. pulvillus, but can be distinguished by the number of adoral membranelles (30–40 vs. 23–27), contractile vacuole (present vs. absent), number of left (27–37 vs. 17–25) and right (27–35 vs. 18–23) marginal cirri, and lepidosomes on the cyst surface (present vs. absent). In a phylogenetic tree, O. seokmoensis is distinctly separated from the O. granulifera clade, but is sister to the Paroxytricha clade. In addition, O. seokmoensis and P. longigranulosa have the smallest genetic difference (d = 0.015, 23 of 1579 nt difference). This close relationship is supported by incomplete dorsal kinety 3 fragmentation and separated macronuclear nodules in resting cysts.     

Taxonomic revision of Chlamydomonas subg. Amphichloris (Volvocales,Chlorophyceae), with resurrection of the genus Dangeardinia and descriptions of Ixipapillifera gen. nov. and Rhysamphichloris gen. nov.

Chlamydomonas (Cd.) is one of the largest but most polyphyletic genera of freshwater unicellular green algae. It consists of 400–600 morphological species and requires taxonomic revision. Toward reclassification, each morphologically defined classical subgenus (or subgroup) should be examined using culture strains. Chlamydomonas subg. Amphichloris is characterized by a central nucleus between two axial pyrenoids, however, the phylogenetic structure of this subgenus has yet to be examined using molecular data. Here, we examined 12 strains including six newly isolated strains, morphologically identified as Chlamydomonas subg. Amphichloris, using 18S rRNA gene phylogeny, light microscopy, and mitochondria fluorescent microscopy. Molecular phylogenetic analyses revealed three independent lineages of the subgenus, separated from the type species of Chlamydomonas, Cd. reinhardtii. These three lineages were further distinguished from each other by light and fluorescent microscopy—in particular by the morphology of the papillae, chloroplast surface, stigmata, and mitochondria—and are here assigned to three genera: Dangeardinia emend., Ixipapillifera gen. nov., and Rhysamphichloris gen. nov. Based on the molecular and morphological data, two to three species were recognized in each genus, including one new species, I. pauromitos. In addition, Cd. deasonii, which was previously assigned to subgroup “Pleiochloris,” was included in the genus Ixipapillifera as I. deasonii comb. nov.     

Sperm storage secretion of female cellar spiders (Pholcus phalangioides; Araneae): a gel-electrophoretic analysis

Glandular secretion in the genital cavity of female Pholcus phalangioides was removed and investigated gel-electrophoretically. Different staining techniques indicate that the secretion contains proteinaceous substances. Some protein fragments show a clear reaction to glyco- and lipoprotein staining procedures.
After copulation, female P. phalangioides keep the spermatozoa embedded in the secretion in the genital cavity. Since the spermatozoa are immobile, i.e. coiled and encapsulated, until shortly before oviposition, a nutritive function of the secretion during sperm storage is unlikely. The viscous quality of the material probably serves to retain the sperm mass in the sperm storage site. Lipoproteinaceous components are assumed to prevent desiccation of the sperm mass. Retaining the spermatozoa and maintaining them in a favourable environment are the most plausible functions of the secretion of female P. phalangioides. In spider species which possess receptacula seminis other functions can be attributed to sperm storage secretions.     

Female genital system of the folding-trapdoor spider Antrodiaetus unicolor (Hentz, 1842) (Antrodiaetidae, Araneae): ultrastructural study of form and function with notes on reproductive biology of spiders

The genitalia of the female folding-trapdoor spider Antrodiaetus unicolor are characterized by two pairs of spermathecae that are arranged in a single row and connected to the roof of the bursa copulatrix. Each single spermatheca is divided into three main parts: stalk, bowl, and bulb, which are surrounded by the spermathecal gland. The epithelium of the spermathecal gland is underlain by a muscle meshwork and consists of different types of cells partly belonging to glandular cell units (Class 3 gland cells) that extend into pores in the cuticle of the stalk and bowl. Interestingly, the bulb lacks glandular pores and is characterized by a weakly sclerotized cuticle. This peculiarly structured bulb probably plays an important role in the discharge of the sperm mass. It is suggested that by contraction of the muscle layer the sperm mass may be squeezed out, when the bulb invaginates and expands into the spermathecal lumen, pushing the sperm to the uterus lumen. Each glandular unit consists of usually one or two central secretory cells that are for the most part surrounded by a connecting cell that again is surrounded by a canal cell. The canal cell, finally, is separated from the other epithelial cells (intercalary cells) located between the glandular units by several thin sheath cells that form the outer enveloping layer of the unit. The secretions are released through a cuticular duct that originates proximally between the apical part of the connecting cell and the apical microvilli of the secretory cells and runs into a pore of the spermathecal cuticle. The glandular products of the Class 3 gland cells likely contribute to the conditions allowing long-term storage of the spermatozoa in this species. Details regarding the ovary, the uterus internus, and the uterus externus are reported. Most of the secretion that composes the chorion of the egg is produced in the ovary. Glandular cell units observed in the uterus externus differ structurally from those in the spermathecae and likely play a different role. Finally, we briefly discuss our results on the female genitalia of A. unicolor in the light of knowledge about the reproductive biology of spiders.     

Balechina and the new genus Cucumeridinium gen. nov. (Dinophyceae), unarmored dinoflagellates with thick cell coverings

The genus Balechina (=subgenus Pachydinium) was established for heterotrophic gymnodinioid dinoflagellates with a thick cell covering. The type species, B. pachydermata (=Gymnodinium pachyderm‐atum), showed numerous fine longitudinal striae, whereas B. coerulea (=G. coeruleum) showed ~24 prominent longitudinal surface ridges or furrows and a distinctive blue pigmentation. We have investigated the morphology and molecular phylogeny of these taxa and the species Gymnodinium cucumis, G. lira and G. amphora from the western Mediterranean, Brazil and Japan. Sudden contractions at the cingulum level were seen in B. pachydermata, which also showed a high morphological variability which included morphotypes that have been described as Amphidinium vasculum, G. amphora, G. dogielii and G. gracile sensu Kofoid and Swezy. Molecular phylogeny based on small subunit rRNA gene sequences revealed that Balechina coerulea, G. cucumis and G. lira formed a clade distantly related to the clade of the type species, B. pachydermata, and G. amphora. We propose the new genus Cucumeridinium for the species with longitudinal ridges and a circular apical groove (Cucumeridinium coeruleum comb. nov., C. lira comb. nov. and C. cucumis comb. nov.), and Gymnodinium canus and G. costatum are considered synonyms of C. coeruleum. The genus Balechina remains for the species with a double‐layer cell covering, bossed surface with fine striae, and an elongated elliptical apical groove. At present, the genus is monotypic containing only B. pachydermata.     

Genital morphology of female goblin spiders (Arachnida: Araneae: Oonopidae) with functional implications

Spider genital morphology usually provides the best characters for taxonomy. Furthermore, functional genital morphology helps to understand the evolution of complex genitalia and their role in the context of sexual selection. The genital systems of most haplogyne spider families are poorly investigated with respect to their morphology. The present study investigates the female genitalia of the oonopids Oonops pulcher, Oonopinus kilikus, and Pseudotriaeris sp. by means of light microscopy and SEM. The male palps are briefly described. Females of O. pulcher store spermatozoa in an anterior and a posterior receptaculum (PRe). The genitalia resemble the primitive dysderoid genitalia supporting the hypothesis that the subfamily Oonopinae contains more basal oonopids. In O. kilikus, the anterior receptaculum is reduced to a sclerite. Spermatozoa are stored in a PRe. The receptacula of Pseudotriaeris sp. are reduced to sclerites. Spermatozoa in the uterus internus indicate that fertilization happens there or in the ovary. The anterior sclerite might serve females to lock the uterus during copulation as suggested for other gamasomorphines. The male palp of O. kilikus is simple, whereas the palps of O. pulcher and Pseudotriaeris sp. appear more complex. Complicated structures on the palp of Pseudotriaeris sp. indicate that males exert copulatory courtship.     

Molecular phylogenetic analysis of <Emphasis Type="Italic">Eugenia</Emphasis> L. (Myrtaceae), with emphasis on southern African taxa

With approximately 1000 species, Eugenia s.str. (excluding Syzygium) is one of the largest genera in the Myrtaceae. Morphological and anatomical studies concluded that two supraspecific groups (X and Y) exist in African members of Eugenia. A multigene approach was used to investigate phylogenetic/molecular relationships within southern African Eugenia, as well as representatives from elsewhere in the world. The cpDNA rpl16 intron data supported the distinctness of species groups X and Y. Little sequence divergence was found within the rpl16 intron of Eugenia. Nuclear sequence data were more variable than cpDNA and also supported the two groups. Combined data enhanced the resolution of closely related species. Species group X is related to Eugenia of the New World whereas the affinity of species group Y lies with Eugenia of the Old World. It is hypothesised that the origin of Eugenia in southern Africa is twofold. The presence of cryptic dioecy (structural androdioecy) in both groups of Eugenia in Africa is probably due to convergence. The suggestion that E. incerta is not native to Africa is supported by molecular evidence. The genus Monimiastrum is embedded within a clade with Mauritian Eugenia.We thank Danielle Florens, J. C. Sevathian and G. dArgent of the Mauritius Herbarium, Reduit, for the collection of all the material from Mauritius; Ed Biffin, CSIRO, Australia, for supplying DNA of E. reinwardtiana and Rhodamnia argentea; A. Salywon, Arizona State Univeristy, Tempe, USA for material from Florida; Timothy Lowrey, University of New Mexico, Albuquerque, USA, for material from Madagascar; and A. Urban, Stefan Neser (ARC, Pretoria) and J. H. Pedrosa-Macedo (Forest Research Foundation, Federal University of Parana, Curitiba, Brazil) for the material from Brazil. The authors are indebted to the National Research Foundation (South Africa) and the University of Pretoria for financial assistance.     

Phylogeny of Habronattus jumping spiders (Araneae: Salticidae), with consideration of genital and courtship evolution

Abstract. DNA sequences from the mitochondrial (including ND1, 16S) and nuclear (EF‐1α) genomes of about ninety‐four species were obtained to reconstruct phylogenetic relationships of Habronattus jumping spiders. Maximum parsimony trees were sought with both separate (mitochondrial, nuclear) and combined analyses; maximum likelihood trees were sought with both separate (ND1, 16S, EF‐1α introns, EF‐1α exons) and combined (mitochondrial, nuclear) analyses. All analyses agreed on some fundamental aspects of the tree, including the monophyly of the previously recognized agilis, amicus, dorotheae and americanus species groups. The deep phylogenetic structure is well resolved, placing the agilis, amicus, tranquillus and dorotheae groups basally. Several other previously unrecognized clades were well supported, including a newly formulated decorus group. The large group of species with modified male first and third legs was supported as monophyletic except for the surprising placement elsewhere of three species of the group. The phenotypic similarities between these three and the others are so detailed and precise that convergence in ornamentation can probably be ruled out. There are hints of phylogenetically distant genetic introgression involving the coecatus group. The combination Habronattus paratus is restored based on the species falling within Habronattus. Regarding patterns of character evolution, there was consistent support for the basal placement of several species groups with a long embolus, suggesting that there were more evolutionary reductions in embolus length than postulated in a previous morphological phylogeny. This is in accord with the expectation that there is a bias to an overly conservative interpretation of a character's evolution if it is interpreted on a phylogeny based in part on that same character. In contrast, the molecular phylogeny did not suggest any instances of the evolutionary transformation of one complex style of courtship into another, a possibility that could have been difficult to detect using the morphological phylogeny because of the same bias to conservativism.