Renewed sampling of inland aquatic habitats in southern Africa yields two novel freshwater crab species (Decapoda: Potamonautidae: Potamonautes)

A recent survey of the freshwater streams of the Mecula and Yao mountains in the Niassa province of Mozambique resulted in the discovery of a new freshwater crab species. This species is genetically and morphologically distinct from described species from Mozambique or its neighbouring countries, and is described as P otamonautes bellarussus sp. nov. In addition, a new semi‐terrestrial burrowing freshwater crab P otamonautes flavusjo sp. nov. from the Highveld of the Mpumalanga province in South Africa is described based on unique genetic and morphological characters. The phylogenetic affinities of the two new species in relation to the described eastern and southern African Potamonautes species is determined and the biogeographic implications are discussed. © 2014 The Linnean Society of London     

Disentangling the divergence and cladogenesis in the freshwater crab species (Potamonautidae: Potamonautes perlatus sensu lato) in the Cape Fold Mountains,South Africa,with the description of two novel cryptic lineages

River networks of major drainages can form barriers that shape the phylogeography of freshwater organisms, particularly those with low dispersal capabilities. Freshwater crab species' distributions can be used to examine hydrological patterns to expose historical drainage interconnectivity. We used molecular sequence data (mitochondrial and nuclear DNA) and divergence time estimations to determine the phylogeography of the freshwater crab, Potamonautes perlatus sensu lato, from six drainage systems along the Cape Fold Mountains, South Africa. Two major clades were detected: clade 1 comprised two geographically discrete haploclades occurring in southern flowing drainages, whereas clade 2 included specimens from western flowing drainages. Divergence time estimations suggested a Pleistocene (c. 2.61 Mya) divergence of P. perlatus s.l. The Pleistocene was associated with arid conditions and drainage contractions. However, it is likely that during the mesic conditions of the Pleistocene, P. perlatus s.l. migrated and diverged into contemporary patterns. We conclude that three lineages are nested within P. perlatus s.l., two representing novel species. Potamonautes perlatus sensu stricto is confined to western flowing drainages. The two novel species both occurring in southern flowing drainages are described here: P otamonautes barbarai sp. nov. occurs in the Gamtoos and Gourits Rivers and P otamonautes barnardi sp. nov. in the Breede River. © 2014 The Linnean Society of London     

Tropical African cedars (Widdringtonia, Cupressaceae): systematics, ecology and conservation status

Phenetic analyses of 24 morphological characters, collected in the field for 150 trees of Widdringtonia in Malawi, Zimbabwe, Mozambique and northern South Africa, show two distinct clusters. One can be identified as W. whytei , a tall, wide-crowned forest tree endemic to Mt Mulanje in Malawi, while the other is W. nodi/bra , a multi-stemmed shrub or narrow-crowned tree, which is common on Mulanje and widespread in southern Africa. These two sympatric species are distinct on phenetic, phylogenetic, ecological and biological grounds. The distinguishing characters of the two species are analysed and discussed and the differences are related to different responses to fire. Widdringtonia whytei is a forest pioneer with limited fire survival ability, while W. nodiflora coppices after fire and is common in fire-prone heathlands. To date it has been assumed that there is a single species of Widdringtonia on Mt Mulanje, and this has led to faulty management and exploitation decisions. Extensive plantations of the economically useless W. nodiflora have been established, in the belief that they are the economically desirable Mulanje cedar, W. whytei , which is now in danger of extinction.     

Genetic structure of mosses Pleurozium schreberi (Willd. ex Brid.) Mitt. and Racomitrium lanuginosum (Hedw.) Brid. along altitude gradients in Hokkaido,Japan

Abstract

The families Pterigynandraceae and Rigodiaceae are reviewed for Africa with reference to the collections made by the BBS Expedition to Mt Mulanje, Malawi in 1991. The single African species of Rigodium (Rigodiaceae) is reported to be quite frequent on Mt Mulanje, and three of the four species of Trachyphyllum (Pterigynandraceae) found in Africa (T. gastrodes, T. dusenii and T. inflexum) are reported, the latter two for the first time from Malawi. On present limited evidence, Malawi seems to be a centre of diversity of the genus.     

Two new cryptic and sympatric species of the king crab parasite Briarosaccus (Cirripedia: Rhizocephala) in the North Pacific

Rhizocephalan barnacles have been reported to parasitize a wide range of king crab species (Lithodidae). So far all these parasites have been assigned to a single species, Briarosaccus callosus Boschma, 1930, which is assumed to have a global distribution. Here we investigate Briarosaccus specimens from three different king crab hosts from the fjord systems of Southeastern Alaska: Lithodes aequispinus Benedict, 1895, Paralithodes camtschaticus (Tilesius, 1815), and Paralithodes platypus (Brandt, 1850). Using molecular markers and by morphological comparison we show that Briarosaccus specimens from these three commercial exploited king crabs are in fact morphologically distinct from B. callosus, and further represent two separate species which we describe. The two new species, Briarosaccus auratum n. sp. and B. regalis n. sp., are cryptic by morphological means and were identified as distinct species by the use of genetic markers (COI and 16S). They occur sympatrically, yet no overlap in king crab hosts occurs, with B. auratum n. sp. only found on L. aequispinus, and B. regalis n. sp. as parasite of the two Paralithodes hosts. © 2015 The Authors. Zoological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London     

A conservation assessment of the freshwater crabs of southern Africa (Brachyura: Potamonautidae)

Recent taxonomic revisions of the freshwater crabs of southern Africa (Angola, Botswana, Lesotho, Mozambique, Namibia, South Africa, Swaziland, Zambia and Zimbabwe) allow accurate depictions of their diversity, distribution patterns and conservation status. The southern African region is home to nineteen species of freshwater crabs all belonging to the genus Potamonautes (family Potamonautidae). These crabs show high levels of species endemism (84%) to the southern African region and to the country of South Africa (74%). The conservation status of each species is assessed using the IUCN (2003) Red List criteria, based on detailed compilations of the majority of known specimens. The results indicate that one species should be considered vulnerable, fifteen species least concern and three species data deficient. The results have been utilized by the IUCN for Red Lists, and may prove useful when developing a conservation strategy for southern Africa’s endemic freshwater crab fauna.     

Ecological Assessment of Two Species of Potamonautid Freshwater Crabs from the Eastern Highlands of Zimbabwe,with Implications for Their Conservation

The spatial ecology of freshwater crabs and their conservation status is largely understudied in Africa. An ecological assessment was conducted at 104 localities in 51 rivers and/or streams in the Eastern Highlands of Zimbabwe whereby the distribution and abundances of freshwater crab species were mapped and the possible drivers of the observed trends in population structure explored. In addition, information on crab utilisation as a food resource by local communities was assessed via face to face interviews across the region. Finally, the conservation status of each species was assessed using the IUCN Red List criteria. Only two crab species Potamonautes mutareensis and Potamonautes unispinus were recorded within the region of study. Potamonautes mutareensis was largely restricted to less impacted environments in the high mountainous river system, whereas P. unispinus was found in low laying areas. In stretches of river where both species were found to co-occur, the species were never sampled from the same site, with P. mutareensis occurring in shallower, faster flowing environments and P. unispinus in deeper, slow flowing sites. Interview results revealed that the local communities, particularly in the southern part of the Eastern Highlands around the Chipinge area, had a considerable level of utilisation (55% of households) on the harvesting of crabs for household consumption during the non-agricultural season (May to September). Results from the IUCN Red List assessment indicate that both species should be considered as “Least Concern”. Threats to freshwater crabs in the Eastern Highlands, however, include widespread anthropogenic impacts such as habitat destruction associated with gold and diamond mining, inorganic and organic pollution and possibly exploitation for human consumption. The current study provides important information and insight towards the possible development of a freshwater crab conservation action plan within the region.     

Presence of itaconic acid in the hemolymph and tissues of the freshwater crab, Potamonautes warreni Calman

Itaconic acid was detected in hemolymph, hepatopancreas, gill and muscle tissue of the freshwater crab, Potamonautes warreni. This metabolite has previously only been found in unicellular organisms and fungi. The measured itaconic acid is either being manufactured by the crab itself or present due to infestation by unicellular organisms or fungi.     

DNA SEQUENCE VARIATION IN THE RIBOSOMAL INTERNAL TRANSCRIBED SPACER REGION OF FRESHWATER CLADOPHORA SPECIES (CHLOROPHYTA)1

Freshwater species of Cladophora (Chlorophyta) are globally distributed and occupy an unusually wide range of ecological habitats. Delineating species is difficult because most easily observed morphological traits are highly variable and because sexual reproduction has not been clearly documented. Synthesizing ecological data on freshwater Cladophora species is problematic because it is unclear whether freshwater Cladophora species comprise many genetically distinct species or a few ecologically and morphologically variable and/or plastic species. We determined nucleotide sequences of the internal transcribed spacer (ITS) region of the nuclear ribosomal cistron of freshwater Cladophora species from a wide range of habitats and geographic locations. We compared these sequences to those derived from culture collections of C. fracta and C. glomerata, the two most commonly reported freshwater Cladophora species. Cladophora fracta and C. glomerata had very similar ITS sequences (95.3%). All other sequences were identical to those from the C. fracta or C. glomerata culture collections with the exception of one California sample that was similar to both C. fracta (95.6%) and C. glomerata (92.4%). ITS genotypes did not correlate with morphology or geography. This analysis shows that common freshwater Cladophora species comprise very few (possibly one) ecologically and morphologically variable species.     

Cryptic diversity in forest shrews of the genus Myosorex from southern Africa,with the description of a new species and comments on Myosorex tenuis

Forest or mouse shrews (Myosorex) represent a small but important radiation of African shrews generally adapted to montane and/or temperate conditions. The status of populations from Zimbabwe, Mozambique, and the north of South Africa has long been unclear because of the variability of traits that have traditionally been ‘diagnostic’ for the currently recognized South African taxa. We report molecular (mitochondrial DNA and nuclear DNA), craniometric, and morphological data from newly collected series of Myosorex from Zimbabwe (East Highlands), Mozambique (Mount Gorogonsa, Gorongosa National Park), and the Limpopo Province of South Africa (Soutpansberg Range) in the context of the available museum collections from southern and eastern Africa and published DNA sequences. Molecular data demonstrate close genetic similarity between populations from Mozambique and Zimbabwe, and this well‐supported clade (herein described as a new species, M yosorex meesteri sp. nov. ) is the sister group of all South African taxa, except for Myosorex longicaudatus Meester & Dippenaar, 1978. Populations of Myosorex in Limpopo Province (herein tentatively assigned to Myosorex cf. tenuis) are cladistically distinct from both Myosorex varius (Smuts, 1832) and Myosorex cafer (Sundevall, 1846), and diverged from M. varius at approximately the same time (2.7 Mya) as M. cafer and Myosorex sclateri Thomas & Schwann, 1905 diverged (2.4 Mya). Morphometric data are mostly discordant with the molecular data. For example, clearly distinct molecular clades overlap considerably in craniometric variables. On the other hand, extreme size differentiation occurs between genetically closely related populations in the Soutpansberg Range, which coincides with the bissection of the mountain range by the dry Sand River Valley, indicating the potential for strong intraspecific phenotypic divergence in these shrews. © 2013 The Linnean Society of London     

Prey preferences in captivity of the freshwater crab Potamonautes lirrangensis from Lake Malawi with special emphasis on molluscivory

Freshwater crabs play an important role for the diversification of shell morphologies in freshwater gastropods. For example, the radiation of the freshwater crab genus Platythelphusa in Lake Tanganyika is thought to have driven shell diversification of the lake’s snail fauna, promoting the evolution of thalassoid shells. No comparable thalassoid snails are known from Lake Malawi. Accordingly, it was hypothesized that the lake’s only freshwater crab, Potamonautes lirrangensis, is not a snail predator. We tested this hypothesis using feeding experiments with specimens caught in the southern part of Lake Malawi. Individual crabs were held in experimental containers offshore and were presented with various food items overnight, after which ingestion frequency was recorded. Potamonautes lirrangensis can be characterized as a scavenger that is opportunistically carnivorous. A preference for fish and snail flesh could be observed, indicating a bias toward carnivory. We observed occasional cracking of the shell in different snail species, with frequent ingestion of artificially crushed specimens, suggesting that crabs do attempt to feed on snails. However, the investigated Lake Malawi gastropods appear to be partly protected against crab predation through thick-walled and low-spired shells (especially Lanistes and Bellamya), obviating the evolution of thalassoid shells carrying rims, ridges, or spines.     

On the marine sister groups of the freshwater crabs (Crustacea: Decapoda: Brachyura)

Freshwater crab sister group relationships with marine eubrachyuran families were investigated. A morphology-based cladistic analysis was conducted on representatives of the freshwater crab families Deckeniidae, Gecarcinucidae, Parathelphusidae, Potamidae, Potamonautidae, Pseudothelphusidae, and Trichodactylidae using a disparate assemblage of marine heterotreme and thoracotreme brachyurans as possible sister groups. The monophyly of the freshwater crabs sensu lato is falsified. The family Trichodactylidae and the marine portunid subfamily Carcininae form basal groups within the superfamily Portunoidea. The monophyly of the Pseudothelphusidae and the Paleotropical freshwater crab families is supported, and this clade is the sister group of the Thoracotremata (Gecarcinidae, Grapsidae s.l., and Ocypodoidea). The origin, groundplan, and diversification of freshwater crabs are discussed in the context of previously published scenarios of their evolution.     

Extensive diversification of pebblesnails (Lithoglyphidae: Fluminicola) in the upper Sacramento River basin, northwestern USA

Mitochondrial DNA sequences from the cytochrome c oxidase subunit I (COI) and cytochrome b (cytb) genes were obtained from the nine extant, previously described species of the northwestern North American freshwater gastropod genus Fluminicola (commonly known as pebblesnails) and from a large number of taxonomically undescribed populations of these animals from the upper Sacramento River basin, California and Oregon, which is composed of the Sacramento River headwaters, and the McCloud and Pit Rivers. Phylogenetic analyses of separate and combined molecular datasets yielded well‐supported and largely congruent trees delineating 13 genetically divergent and morphologically distinctive upper Sacramento basin lineages, which we describe as new species. These include two groups of closely related and geographically proximal species that are further united by unique radular or shell features. Most of these novelties have narrow geographical distributions and are restricted to headspring areas, whereas several are more wide ranging and typically occupy larger, well‐integrated habitats. The highly endemic fauna of upper Sacramento River pebblesnails is not a single species flock, but instead a polyphyletic assemblage spread among four separate clades. Our phylogeny, together with the application of a COI molecular clock for Fluminicola, suggests that upper Sacramento River clades originated as a result of late Neogene separation of this basin from neighbouring regions (northwestern Great Basin, Klamath River basin), which is consistent with previous biogeographical hypotheses based on the distributions of fishes. The upper Sacramento River pebblesnails evolved in association with the complex late Cenozoic history of regional landscape and drainage and diversification was also facilitated by the invasion of and adaptation to insular spring habitats. Our findings are consistent with the generally limited dispersal ability and geologically ancient (mid‐Tertiary) age of this genus and imply that other portions of northwestern North America may also harbour a large number of undescribed pebblesnail species. Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149 , 371–422. No claim to original US government works.     

Ultrastructure of spermatozoa and spermatophores of old world freshwater crabs (Brachyura: Potamoidea: Gecarcinucidae,Potamidae, and Potamonautidae)

We investigated the ultrastructure of spermatozoa and spermatophores of 19 palaeotropical freshwater crab species [12 species of the Gecarcinucidae, 6 of the Potamidae (Potamiscinae), and 1 species of the Potamonautidae (Deckeniinae: Hydrothelphusini)]. The investigated Potamiscinae have densely packed coenospermic spermatophores with the exception of Thaiphusa sirikit and Johora singaporensis that exhibit cleistospermia. In contrast, in the Gecarcinucidae the spermatozoa are loosely embedded in a mucous matrix. The gecarcinucid and potamiscine sperm differ, furthermore, in acrosomal structure and size. The acrosome in the Gecarcinucidae is much smaller and spherical, while the larger acrosome in the Potamiscinae has the tendency to be depressed. In the Potamiscinae, an additional middle acrosomal zone evolved between the acrosome ray zone and the outer acrosomal zone. Within the Gecarcinucidae, a differentiation into two groups (Gecarcinucinae and Parathelphusinae) is not supported by the present spermatological data. The sperm morphology of Hydrothelphusa aff. madagascariensis (Potamonautidae: Deckeniinae) differs from Potamonautes sidneyi (Potamonautidae: Potamonautinae) in acrosomal size and shape, and in the absence of a periopercular rim. A closer relationship of Deckeniinae and Gecarcinucidae cannot be confirmed by spermatology. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Genetic and morphological differentiation of the mangrove crab Perisesarma guttatum (Brachyura: Sesarmidae) along an East African latitudinal gradient

The genetic structure and morphometric differentiation of mangrove crab Perisesarma guttatum populations were examined among shelf connected locations along a latitudinal gradient on the East African coast. Over 2200 specimens were sampled from 23 mangrove sites for geometric morphometrics analysis. Population genetic analyses of mitochondrial cytochrome c oxidase subunit I (COI) DNA sequences were used to evaluate connectivity among populations. A total of 73 haplotypes were detected, and almost no haplotypes were found in common between two highly supported phylogeographic clades: southern Mozambique (Inhaca Island and Maputo Bay) and a northern clade that included north Mozambique, Tanzania and Kenya. These two clades were identified based on the species' populations pairwise genetic differentiation and geographical location. Φ_ST values were considerably high between the two clades, indicating the presence of significant population genetic structure between Kenya and South Mozambique. However, each clade was composed of genetically similar populations along the latitudinal gradient, and no significant population structure was found within each clade because the Φ_st values were not significant. The morphometric analysis corroborated the division into two clades (i.e. Inhaca Island/Maputo Bay and northern populations) and also detected less shape variation among populations that were few kilometres apart. The significant spatial genetic structuring between the southern and the northern populations of P. guttatum along the geographic gradient under study, combined with morphological differences, suggests that these populations may be considered as cryptic species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 28–46.     

Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago

Gigantism and dwarfism evolve in vertebrates restricted to islands. We describe four new species in the Rhinolophus hildebrandtii species-complex of horseshoe bats, whose evolution has entailed adaptive shifts in body size. We postulate that vicissitudes of palaeoenvironments resulted in gigantism and dwarfism in habitat islands fragmented across eastern and southern Africa. Mitochondrial and nuclear DNA sequences recovered two clades of R. hildebrandtii senso lato which are paraphyletic with respect to a third lineage (R. eloquens). Lineages differ by 7.7 to 9.0% in cytochrome b sequences. Clade 1 includes R. hildebrandtii sensu stricto from the east African highlands and three additional vicariants that speciated across an Afromontane archipelago through the Plio-Pleistocene, extending from the Kenyan Highlands through the Eastern Arc, northern Mozambique and the Zambezi Escarpment to the eastern Great Escarpment of South Africa. Clade 2 comprises one species confined to lowland savanna habitats (Mozambique and Zimbabwe). A third clade comprises R. eloquens from East Africa. Speciation within Clade 1 is associated with fixed differences in echolocation call frequency, and cranial shape and size in populations isolated since the late Pliocene (ca 3.74 Mya). Relative to the intermediate-sized savanna population (Clade 2), these island-populations within Clade 1 are characterised by either gigantism (South African eastern Great Escarpment and Mts Mabu and Inago in Mozambique) or dwarfism (Lutope-Ngolangola Gorge, Zimbabwe and Soutpansberg Mountains, South Africa). Sympatry between divergent clades (Clade 1 and Clade 2) at Lutope-Ngolangola Gorge (NW Zimbabwe) is attributed to recent range expansions. We propose an “Allometric Speciation Hypothesis”, which attributes the evolution of this species complex of bats to divergence in constant frequency (CF) sonar calls. The origin of species-specific peak frequencies (overall range = 32 to 46 kHz) represents the allometric effect of adaptive divergence in skull size, represented in the evolution of gigantism and dwarfism in habitat islands.     

European cave shrimp species (Decapoda: Caridea: Atyidae), redefined after a phylogenetic study; redefinition of some taxa,a new genus and four new Troglocaris species

Endemic atyids of southern Europe have been ascribed to Dugastella, and to subterranean Typhlatya and Troglocaris: Dugastella is epigean, and Typhlatya and Troglocaris are subterranean. An extensive collection from all centres of distribution in southern Europe (excepting the Caucasus) was morphologically examined. A taxonomic redefinition of the group, at different levels, is based on recently published and newly generated molecular phylogeny, whereas newly established taxa have also been morphologically defined. The accordance between the phylogenetic tree and the geographical distributions suggested that a re‐evaluation of some traditionally used morphological characters should generate the most parsimonious solution: this enabled a novel taxonomic division. Gallocaris gen. nov. is erected for the French Troglocaris inermis Fage, 1937, which is more closely related to the epigean Dugastella valentina (Ferrer Galdiano, 1924) than to its supposed congeners. Both western Mediterranean Typhlatya species are closely related to their Caribbean congeners. All other European cave shrimps constitute a monophylum, Troglocaris, which is divisible into subgenera (already with available names): the holo‐Dinaric Troglocaris (Troglocaris) Dormitzer, 1853, south‐eastern mero‐Dinaric Troglocaris (Troglocaridella) Babi?, 1922, and Troglocaris (Spelaeocaris) Matja?i?, 1956, and the Caucasian Troglocaris (Xiphocaridinella) Sadovsky, 1930. Four new species are described: Troglocaris (Troglocaris) bosnica sp. nov. , Troglocaris (Spelaeocaris) prasence sp. nov. , Troglocaris (Spelaeocaris) kapelana sp. nov. , and Troglocaris (Spelaeocaris) neglecta sp. nov. The distribution of all established species is shown, and the value of the morphological characters is discussed. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 786–818.     

At the foot of the shrew: manus morphology distinguishes closely‐related Cryptotis goodwini and Cryptotis griseoventris (Mammalia: Soricidae) in Central America

Small‐eared shrews (Mammalia, Soricidae) of the New World genus Cryptotis are distributed from eastern North America to the northern Andes of South America. One well‐defined clade in this genus is the Central American Cryptotis mexicana group, whose members are set off from other species in the genus by their variably broader fore feet and more elongate and broadened fore claws. Two species in the C. mexicana group, Cryptotis goodwini Jackson and Cryptotis griseoventris Jackson, inhabit highlands in Guatemala and southern Mexico and are presumed to be sister species whose primary distinguishing feature is the larger body size of C. goodwini. To better characterize these species and confirm the identification of recently‐collected specimens, we obtained digital X‐ray images of the manus from large series of dried skins of both species. Measurements of the metacarpals and phalanges successfully separated most specimens of C. goodwini and C. griseoventris. These measurements also show that the fore feet of C. griseoventris from Chiapas, Mexico, are morphologically distinct from those of members of the species inhabiting Guatemala. Univariate, bivariate, and multivariate analyses indicate that fore foot characters are more conservative within species of the C. mexicana group than are cranio‐mandibular characters. Patterns of evolution of fore foot characters that superficially appear to be linear gradations are actually more complex, illustrating individual evolutionary trajectories. No claim to original US government works. Journal compilation © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 118–134.