David Andrew Barraclough

Is Millipede Taxonomy Based on Gonopod Morphology Too Inclusive? Observations on Genetic Variation and Cryptic Speciation in Bicoxidens flavicollis (Diplopoda: Spirostreptida: Spirostreptidae)

ABSTRACT The structure of the male gonopods of millipedes has been considered to be species-specific. As such, gonopods—which aid in copulation and sperm transfer—are used in the taxonomic diagnosis and description of species. However, it was recently demonstrated that gonopod morphology is not always characteristic of species. Diagnoses based on gonopod morphology can therefore result in underestimation of taxonomic diversity amongst millipedes. On the basis of this observation, we examined genetic variation in two populations (approximately 250 km apart) of a widely distributed and colour-polymorphic southern African millipede, namely Bicoxidens flavicollis Attems, 1928. An analysis of genetic divergence based on 520 nucleotides of the mitochondrial cytochrome oxidase 1 gene, and 684 nucleotides of the mitochondrial 16S rRNA gene, demonstrated high levels of divergence (19.09% for cytochrome oxidase 1 and 6.66% for 16S rRNA) between the two populations. These results suggest the presence of cryptic species in B. flavicollis and, furthermore, corroborate observations that taxonomy based on gonopod morphology may be too inclusive.

The South African keystone pollinator Moegistorhynchus longirostris (Wiedemann, 1819) (Diptera: Nemestrinidae): notes on biology, biogeography and proboscis length variation

ABSTRACT Moegistorhynchus longirostris Wiedemann, 1819 has the longest proboscis relative to body size of all known insects. It is a keystone species along the west coast of South Africa, where it pollinates, partly or exclusively, the long-tubed flowers of at least 20 species of Iridaceae, Geraniaceae and Orchidaceae. M. longirostris has been widely discussed in the pollination biology literature and has a suggested co-evolutionary arms race between it and the long-tubed flowers it pollinates. A current taxonomic revision has presented new biogeographical and morphometric data. Available records suggest that the species is widely distributed along the west coast of South Africa between about 29°30′S and 34°15′S, a distance of almost 500 km. It is restricted to sandy, lowland coastal plains at less than 300 m, where it occurs in nine different vegetation types across seven bioregions and is consequently not a habitat specialist. Proboscis length from 13 localities ranged from 32 to 83 mm, and mean proboscis length at these localities from 32 to 71 mm. There was a decline in proboscis length with increasing latitude south. Research investigating a possible allometric relationship between body size and proboscis length, and between latitude and proboscis length, is required.

Do cytochrome c oxidase 1 Gene Sequences Differentiate Species of Spirostreptid Millipedes (Diplopoda: Spirostreptida: Spirostreptidae)?

ABSTRACT The aim of this study was to use cytochrome c oxidase 1 (CO1) sequences to recover a phylogeny for seven morphologically described spirostreptid millipede taxa from southern Africa, and to evaluate the correspondence between morphological and molecular phylogenies. Genetic p-distance generally increased with taxonomic divergence: inter-specific mean 15.33 % (14.09 % –17.02 %), inter-generic mean 18.43 % (6.83 %–26.81 %) and inter-order mean 24.16 % (range 18.56 %–30.77 %). Congruent Bayesian, maximum parsimony and neighbour-joining analyses of 520 nucleotides of the CO1 gene resolved the orders Spirostreptida, Julida and Callipodida. Members of genera within the Spirostreptidae (Archispirostreptus, Bicoxidens, Cacuminostreptus, Doratogonus, Orthoporoides, Plagiotaphrus and Spirostreptus) formed a single clade within which a sample of Thyropygus (family Harpagophoridae) was paraphyletically nested. Phylogenetic analyses failed to recover support for the genera Doratogonus, Bicoxidens, Archispirostreptus and Spirostreptus, as representatives of these genera were not monophyletic. Samples morphologically identified as the same species (Bicoxidens flavicollis) were part of two different clades, one of which was well supported and otherwise contained members of Doratogonus. This high level of divergence (mean 12.64 %) between morphologically identified spirostreptid millipede sister species could indicate that changes in genital morphology occur rather slowly relative to CO1 sequence substitution, and may underestimate species diversity.

Description of Two New Species of Zinophora Chamberlin, 1927 (Diplopoda: Spirostreptida: Harpagophoridae), with Discussion of Species Groups in the Genus

ABSTRACT The millipede genus Zinophora (Chamberlin, 1927) was previously revised based on the species present south of the Zambezi and Kunene Rivers. Since that revision, two additional new species have been discovered and are here described: Zinophora lobata (Mozambique) and Z. taromberai (Zimbabwe and Tanzania). This brings the total number of described species in the genus to 21. An updated key to species in the genus is presented. Detailed illustrations of the gonopods supplement the descriptions and a distribution map of Z. taromberai in Zimbabwe is provided. There appear to be up to three species groups in the genus and the character states defining these groups are discussed.

A striking new species of Prosoeca Schiner, 1867 (Diptera: Nemestrinidae): An important pollinator from the Bokkeveld Plateau, Northern Cape Province, South Africa

For more than 20 years an undescribed species of Prosoeca has been referred to in numerous publications by pollination biologists, evolutionary biologists and ecologists, originally as being part of the Prosoeca peringueyi Lichtwardt, 1920, pollination guild. Ongoing research in these and related fields has necessitated the formal description of this large-bodied, striking new species, with a proboscis 1.5-2.3 x body length (mean proboscis length ± SD 36.25 ± 3.90 mm). Prosoeca marinusi Barraclough sp. nov. is described from a long series from the Hantam National Botanical Garden, Nieuwoudtville, Bokkeveld Plateau, Northern Cape Province, South Africa. The species is a narrow-range endemic and is restricted to the Nieuwoudtville area. Observations on the biology of the species are also presented. Prosoeca marinusi Barraclough sp. nov. is the only or main pollinator of at least four plant species (all regional endemics) in the family Iridaceae that flower from August to September. During August, Lapeirousia oreogena Goldblatt and Babiana vanzyliae L. Bolus are the main host flowers, while later in the spring season, Babiana framesii L. Bolus is most abundant in the Nieuwoudtville area.

Modified fore Femora in a New Species of Afrocamilla Barraclough: A Morphological Innovation in the Camillidae (Diptera)

ABSTRACT A new species of Afrocamilla is described from a series of females from the Bale Mountains in southeastern Ethiopia. The new species is unique within the Camillidae in having markedly swollen fore femora with a posteroventral protrusion of cuticle bearing a dagger-like spine on each femur. The ornamentation of the fore femora of other Camillidae is reviewed. Their possible functions in A. femorata sp. n. are evaluated, and based on the position of the spines it is suggested that they may be used to assist will) movement over uneven substrates or for breaking open dung pellets for egg deposition.