Nucleotide sequence variation in the mitochondrial 12S rRNA gene and the phylogeny of African mole-rats (Rodentia: Bathyergidae).

Mitochondrial DNA (mtDNA) sequence variation was examined in eight taxa of the African rodent family Bathyergidae, as well as in two taxa representative of the Old-World hystricognathid rodent families Petromyidae and Thryonomyidae. A total of 812 bp, constituting domains I-III of the 12S ribosomal rRNA gene, were compared for each taxon. The high levels of intrafamilial mtDNA sequence divergence observed (average 16.8, range 3.5-23.2) support an ancient origin for the five genera, 20-38 Mya. These data do not support the current subfamilial groupings of the Bathyergidae. The eastern African naked mole-rat, Heterocephalus glaber, is the most basal representative of the family, with the silvery mole-rat, Heliophobius, being the next most basal. South African forms [dune, common, and cape mole-rats (Bathyergus, Cryptomys, and Georychus, respectively)] group together. The independent origin of the common mole-rat, relative to the naked mole-rat, suggests that complex social systems evolved in parallel along different bathyergid lineages. The 12S rRNA gene is not evolving at a higher rate within the rodent lineages, relative to that seen for artiodactyls and primates. Bathyergid rodents appear to fall at an extreme end of the spectrum of mammalian variation, with respect to both transition/transversion ratios and divergence, showing much lower transition/transversion ratios than those previously reported for intrafamilial comparisons.     

Growth and development in six species of African mole-rats (Rodentia: Bathyergidae)

The bathyergid mole-rats provide a unique example of a family of subterranean rodents exhibiting a broad spectrum of sociality. Three genera comprise solitary, strongly territorial individuals whereas two genera are social. This sociality culminates in the eusocial naked mole-rat, Heterocephalus glaber . The pups of solitary mole-rats disperse, establish and thereafter defend their own burrow systems when approximately two months old, whereas those of social genera join an established natal colony. This paper examines whether these different lifestyles are reflected in the early development and rate of growth of pups of mole-rats.
Although the trends are not clear-cut, it is apparent that the pups of solitary genera grow and mature more rapidly than those from social genera. Thus, the growth rate constant ( K ) for the first70–80 days of postnatal growth (using the Gompertz equation) for the solitary genera was between 0.042 and 0.052 day⁻¹, whereas that of the social mole-rats was considerably lower (0.01 5 day⁻¹). Similarly the mean growth rates of solitary genera ranged between 3.3 and 1.227g/day while those of the social mole-rats were 0.229-0.233 g/day.
The pattern of development and the rates of growth in solitary bathyergids are similar to those of other solitary subterranean rodents. One interesting feature common to all the social genera studied to date was that the first pups recruited to a 'new colony', consisting of a reproductive pair of adult mole-rats, grew at a significantly faster rate than pups born to an established colony.     

Phylogeographical patterns of genetic divergence and speciation in African mole-rats (Family: Bathyergidae)

African mole-rats are subterranean Hystricomorph rodents, distributed widely throughout sub-Saharan Africa, and displaying a range of social and reproductive strategies from solitary dwelling to the 'insect-like' sociality of the naked mole-rat, Heterocephalus glaber. Both molecular systematic studies of Rodentia and the fossil record of bathyergids indicate an ancient origin for the family. This study uses an extensive molecular phylogeny and mitochondrial cytochrome b and 12s rRNA molecular clocks to examine in detail the divergence times, and patterns of speciation of the five extant genera in the context of rift valley formation in Africa. Based on a value of 40-48 million years ago (Myr) for the basal divergence of the family (Heterocephalus), we estimate divergence times of 32-40 Myr for Heliophobius, 20-26 Myr for Georychus/Bathyergus and 12-17 Myr for Cryptomys, the most speciose genus. While early divergences may have been independent of rifting, patterns of distribution of later lineages may have been influenced directly by physical barriers imposed by the formation of the Kenya and Western Rift, and indirectly by accompanying climatic and vegetative changes. Rates of chromosomal evolution and speciation appear to vary markedly within the family. In particular, the genus Cryptomys appears to have undergone an extensive radiation and shows the widest geographical distribution. Of the two distinct clades within this genus, one exhibits considerable karyotypic variation while the other does not, despite comparatively high levels of sequence divergence between some taxa. These different patterns of speciation observed both within the family and within the genus Cryptomys may have been a result of environmental changes associated with rifting.     

Circadian rhythms of locomotor activity in solitary and social species of African mole-rats (family: Bathyergidae)

Mole-rats are strictly subterranean and hardly, if ever, come into contact with external light. As a result, their classical visual system is severely regressed and the circadian system proportionally expanded. The family Bathyergidae presents a unique opportunity to study the circadian system in the absence of the classical visual system in a range of species. Daily patterns of activity were studied in the laboratory under constant temperature but variable lighting regimes in individually housed animals from 3 species of mole-rat exhibiting markedly different degrees of sociality. All 3 species possessed individuals that exhibited endogenous circadian rhythms under constant darkness that entrained to a light-dark cycle. In the solitary species, Georychus capensis, 9 animals exhibited greater activity during the dark phase of the light cycle, while 2 individuals expressed more activity in the light phase of the light cycle. In the social, Cryptomys hottentotus pretoriae, 5 animals displayed the majority of their activity during the dark phase of the light cycle and the remaining 2 exhibited more activity during the light phase of the light cycle. Finally in the eusocial Cryptomys damarensis, 6 animals displayed more activity during the light phase of the light cycle, and the other 2 animals displayed more activity during the dark phase of the light cycle. Since all three mole-rat species are able to entrain their locomotor activity to an external light source, light must reach the SCN, suggesting a functional circadian clock. In comparison to the solitary species, the 2 social species display a markedly poorer response to light in all aspects. Thus, in parallel with the sociality continuum, there exists a continuum of sensitivity of the circadian clock to light.     

A tangled history: patterns of major histocompatibility complex evolution in the African mole-rats (Family: Bathyergidae)

Phylogenetic trees based upon major histocompatibility complex (MHC) gene sequences, particularly those encompassing sites encoding the antigen recognition site, are often discordant with the species tree. It has been argued that the principal cause of such discordance is the presence of ancestrally derived polymorphisms persisting through speciation events as a consequence of selection. In the present study, we examine the evolution of the MHC class II DQα1 gene in an unusual family of hystricomorph rodents, the African mole-rats (Family: Bathyergidae). We show that there is a high level of trans-species polymorphism and that this is a result of positive selection. Furthermore, the major lineages of the gene tree are characterized by allelic motifs occurring in regions that coincide with the pocket domains of the putative antigen recognition site, a region that has been shown to be under positive selection in a number of MHC genes from a range of species. Finally, these alleles may have been retained for at least 48 million years. This is significantly older than the estimate for the equivalent primate locus and appears to be one of the oldest documented sets of MHC alleles. We suggest that these allelic motifs possess polymorphisms that have been immunologically important to African mole-rats over long periods of evolutionary history.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 493–503.     

Thermal biology of a strictly subterranean mammalian family,the African mole-rats (Bathyergidae,Rodentia) - a review

African mole-rats are subterranean rodents, which rarely if ever leave the safety of their burrow systems. The environment of the burrows is humid, with relatively stable temperatures, and may have a hypoxic and hypercapnic atmosphere. One of crucial problems related to the subterranean way of life in mammals is avoidance of overheating, because traditional mammalian cooling mechanisms are not effective under high humidity. In African mole-rats, a variety of adaptations have evolved in response to this and other challenges of the underground ecotope. Traditionally, attention has been devoted mainly to the naked mole-rat Heterocephalus glaber, which became popular as a result of its eusociality and absence of fur, both being unique phenomena in small mammals. Despite more recent research, information on other species is still relatively limited and patchy. I review the results of studies on African mole-rats that are relevant for the understanding of their energetics and thermal biology. Attention is paid to the parameters of the burrow environment, which represent the main selection pressures shaping their physiology. In addition, an overview is given of the morphological, physiological and behavioural adaptations helping mole-rats to face temperature extremes, mechanisms by which they deal with a surplus of metabolic heat and how changes in ambient temperature influence their daily activity. The naked mole-rat is compared to its furred relatives to determine whether this species is really exceptional from the point of thermal biology. An ordination analysis was conducted using published data on mole-rat body temperature, thermoneutral zone, resting metabolic rate and thermal conductance. Most of the variability in these characteristics was found to be explained by body mass, followed by temperature characteristics of climate, but not precipitation, of the species distributional ranges. This analysis shows that the naked mole-rat is comparable to the other mole-rat species in these physiological characteristics.     

Hind foot drumming: Morphofunctional analysis of the hind limb osteology in three species of African mole-rats (Bathyergidae)

Hind foot drumming is a form of seismic signaling that plays a vital role in the communication of several Bathyergidae species. Hind foot drumming is initiated by the rapid movement of the whole hind limb by flexion and extension of the hip and knee. This study aimed to determine if morphological adaptations of the hind limb osteology were measurable using established morphometric analyses in two drumming (Bathyergus suillus and Georychus capensis) and one non-drumming (Cryptomys hottentotus natalensis) African mole-rat species. Forty-three linear measurements of the hind limb were taken in 48 limbs (n = 16 limbs per species) and 32 indices were calculated. Mixed model analysis of variance was used to compare the three species and sexes within a species. Thirteen indices had significant differences between species. Eleven indices had significant differences between sexes within a species. Significant differences between the drumming (B. suillus and G. capensis) and the non-drumming species were observed in three indices. The femoral greater trochanter was relatively shorter in the drumming species compared to the non-drumming species, which is proposed to allow for increased hip joint mobility, thereby permitting drummers to move their limbs at the rapid speed required to generate seismic signals. Furthermore, the small in-lever (shorter greater trochanter) may increase the velocity of limb motion. The robust tibias in the drumming species, as indicated by the tibial robustness index, are likely to counter the additional biomechanical load caused by the muscles involved in hind foot drumming. The relatively small hind feet seen in the drumming species allows for reduced limb weight needed for the rapid extension and flexion motion required during hind foot drumming. The significant differences reflected in the hind limb osteological indices between B. suillus and G. capensis and the non-drumming species are indicative of adaptations for hind foot drumming.     

Coefficients of digestibility and nutritional values of geophytes and tubers eaten by southern African mole-rats (Rodentia: Bathyergidae)

Subterranean rodents have high energy requirements when they are excavating their burrows. This study investigates the energy available to, and the efficiency with which it can be extracted by, four species of bathyergid mole-rats fed natural diets ranging from the underground storage organs of geophytes to grass roots and leaves.
The digestibility coefficients of geophytes ranged from 53% for the fibrous tuber of the gemsbok cucumber to 95–7% for corms and bulbs. One species, Bathyergus suillus whose diet consists of over 80% grass, had a digestibility coefficient of 87% on an all grass diet.
All species had similar coefficients of digestibility of > 90% when fed on a uniform diet of sweet potato.
Bulbs and corms had a low fibre content (3–3–4%), high calorific value (15–16kJ/g) and high digestibility coefficients (95–7–96%) and on this diet the mole-rats maintained their body mass. Food of lower digestibility tended to have a high fibre content (8–2–45%) and, with the exception of B. suillus , although the mole-rats consumed a greater quantity of food, they lost mass. The sweet potato had a low fibre content (4–1 %) but was energetically very similar to bulbs and corms (15–5 kJ/g).
Geophytes which have low fibre contents are generally small (1–20 g), whereas geophytes with high fibre contents are much larger (30–2000 g) and often occur in more arid zones.     

Cytochrome b sequence analysis reveals differential molecular evolution in African mole-rats of the chromosomally hyperdiverse genus Fukomys (Bathyergidae, Rodentia) from the Zambezian region

African mole-rats (Bathyergidae, Rodentia) of the (eu)social genus Fukomys are one of the most speciose mammal genera endemic to Sub-Saharan Africa. Fukomys distributed in the Zambezian phytochorion is characterized by extreme chromosomal variation (2n=40-78). We inferred a molecular phylogeny of Zambezian Fukomys to resolve the interrelationships and the evolutionary history of the known chromosomal races. We sequenced the entire cytochrome b gene (1140bp) for a total of 66 specimens representing 18 karyotypical races from Zambia. An additional 31 sequences were retrieved from GenBank including data on all other chromosomal races. The haplotypes belonging to a small chromosomal race from Salujinga cluster with the Fukomys mechowii (Giant mole-rat) haplotypes. Differential degrees of chromosomal variation are observed among the major mole-rat clades, which is most pertinent when comparing the central Zambezian Fukomys micklemi and the northern Zambezian Fukomys whytei clades. The karyotypically hyper-diverse (12 known chromosomal races) Fukomys micklemi clade shows low levels of cytochrome b sequence divergence. Within the F. whytei clade we find a more conservative pattern of chromosomal diversification (three known chromosomal races) while the levels of sequence divergence are much higher then in the F. micklemi clade. Our results suggest that chromosomal changes may drive phyletic divergence and, eventually, speciation. The observed cladogenetic events during the Plio-Pleistocene within the F. mechowii, F. whytei, F. damarensis and F. micklemi clades appear to coincide with climatically mediated speciation bursts in other savannah dwelling mammals, including hominids. Based on the molecular data presented, combined with morphological and chromosomal data, the taxonomic implication seems to be that Fukomys may contain several (undescribed) cryptic species.     

A phylogeny of cycads (Cycadales) inferred from chloroplast matK gene, trnK intron, and nuclear rDNA ITS region

Phylogenetic relationships among the three families and 12 living genera of cycads were reconstructed by distance and parsimony criteria using three markers: the chloroplast matK gene, the chloroplast trnK intron and the nuclear ITS/5.8S rDNA sequence. All datasets indicate that Cycadaceae (including only the genus Cycas) is remotely related to other cycads, in which Dioon was resolved as the basal-most clade, followed by Bowenia and a clade containing the remaining nine genera. Encephalartos and Lepidozamia are closer to each other than to Macrozamia. The African genus Stangeria is embedded within the New World subfamily Zamiodeae. Therefore, Bowenia is an unlikely sister to Stangeria, contrary to the view that they form the Stangeriaceae. The generic status of Dyerocycas and Chigua is unsupportable as they are paraphyletic with Cycas and the Zamia, respectively. Nonsense mutations in the matK gene and indels in the other two datasets lend evidence to reinforce the above conclusions. According to the phylogenies, the past geography of the genera of cycads and the evolution of character states are hypothesized and discussed. Within the suborder Zamiieae, Stangeria, and the tribe Zamieae evolved significantly faster than other genera. The matK gene and ITS/5.8S region contain more useful information than the trnK intron in addressing phylogeny. Redelimitations of Zamiaceae, Stangeriaceae, subfamily Encephalartoideae and subtribe Macrozamiineae are necessary.     

Patterns of MHC selection in African mole-rats, family Bathyergidae: the effects of sociality and habitat

African mole-rats are a family of rodents exhibiting an eclectic range of social behaviour and occupying a variety of habitat types. These differences are likely to impact upon the risk of parasite transmission and virulence, with increasing sociality predicted to correspond to an increased risk of transmission. We investigate these factors by analysing the major histocompatibility complex (MHC), a set of genes responsible for encoding highly variable intermediaries of the vertebrate adaptive immune response. To this end we assessed selection at exons 2 and 3 of the MHC class II DQalpha1 gene of four African mole-rat species representing a range of social behaviours. We demonstrate that: (i) the overall pattern of selection at these exons differentiates according to the predicted function of different regions, with the presence of positive selection indicating the likely influence of host-parasite coevolution; and (ii) contrary to the often observed and predicted positive correspondence between sociality and the risk of parasite transmission, two highly social African mole-rat species in fact appear to have comparatively weak positive selection, suggesting diminished host immunity and thus a low overall risk of parasite transmission.     

Energetics in a solitary subterranean rodent, the silvery mole-rat, Heliophobius argenteocinereus, and allometry of RMR in African mole-rats (Bathyergidae)

Low resting metabolic rate (RMR) in subterranean rodents used to be considered as a physiological adaptation to cope with stresses of the belowground environment. In African mole-rats (Bathyergidae, Rodentia), RMR was reported to be independent of body mass. This deviation from a general mammalian pattern was considered a precondition for evolution of eusociality, occurring in some bathyergids. We measured metabolic rate and thermoregulation in the silvery mole-rat, Heliophobius argenteocinereus, the only bathyergid genus for which well-supported, comparable data were still missing. Low RMR (154.04 mL O(2) h(-1), which is 82% of the value predicted for a rodent) corresponds to the value expected in a subterranean rodent. Broad range of the thermoneutral zone (25-33 degrees C) and only slightly higher conductance (17.3 mL O(2) h(-1) degrees C(-1), i.e. 112.5% of that predicted for subterranean mammals) indicate that H. argenteocinereus is adapted to lower burrow temperatures rather than to high temperatures. Low RMR in this solitary species, as in other subterranean rodents in general, is probably associated particularly with high energetic cost of foraging. Our results combined with data on other mole-rats show clearly that RMR within the Bathyergidae is mass-dependent.     

Molecular phylogeny of Pemphiginae (Hemiptera: Aphididae) inferred from nuclear gene EF-1alpha sequences

Three traditional tribes of Fordini, Pemphigini and Eriosomatini comprise Pemphiginae, and there are two subtribes in Fordini and Pemphigini, respectively. Most of the species in this subfamily live heteroecious holocyclic lives with distinct primary host specificity. The three tribes of Pemphigini (except Prociphilina), Eriosomatini and Fordini use three families of plants, Salicaceae (Populus), Ulmaceae (Ulums) and Anacardiaceae (Pistacia and Rhus), as primary hosts, respectively, and form galls on them. Therefore, the Pemphigids are well known as gall makers, and their galls can be divided into true galls and pseudo-galls in type. We performed the first molecular phylogenetic study of Pemphiginae based on molecular data (EF-1alpha sequences). Results show that Pemphiginae is probably not a monophylum, but the monophyly of Fordini is supported robustly. The monophyly of Pemphigini is not supported, and two subtribes in it, Pemphigina and Prociphilina, are suggested to be raised to tribal level, equal with Fordini and Eriosomatini. The molecular phylogenetic analysis does not show definite relationships among the four tribes of Pemphiginae, as in the previous phylogenetic study based on morphology. It seems that the four tribes radiated at nearly the same time and then evolved independently. Based on this, we can speculate that galls originated independently four times in the four tribes, and there is no evidence to support that true galls are preceded by pseudo-galls, as in the case of thrips and willow sawflies.     

Extended longevity of reproductives appears to be common in Fukomys mole-rats (Rodentia, Bathyergidae)

African mole-rats (Bathyergidae, Rodentia) contain several social, cooperatively breeding species with low extrinsic mortality and unusually high longevity. All social bathyergids live in multigenerational families where reproduction is skewed towards a few breeding individuals. Most of their offspring remain as reproductively inactive "helpers" in their natal families, often for several years. This "reproductive subdivision" of mole-rat societies might be of interest for ageing research, as in at least one social bathyergid (Ansell's mole-rats Fukomys anselli), breeders have been shown to age significantly slower than non-breeders. These animals thus provide excellent conditions for studying the epigenetics of senescence by comparing divergent longevities within the same genotypes without the inescapable short-comings of inter-species comparisons. It has been claimed that many if not all social mole-rat species may have evolved similar ageing patterns, too. However, this remains unclear on account of the scarcity of reliable datasets on the subject. We therefore analyzed a 20-year breeding record of Giant mole-rats Fukomys mechowii, another social bathyergid species. We found that breeders indeed lived significantly longer than helpers (ca. 1.5-2.2fold depending on the sex), irrespective of social rank or other potentially confounding factors. Considering the phylogenetic positions of F. mechowii and F. anselli and unpublished data on a third Fukomys-species (F. damarensis) showing essentially the same pattern, it seems probable that the reversal of the classic trade-off between somatic maintenance and sexual reproduction is characteristic of the whole genus and hence of the vast majority of social mole-rats.     

Chloroplastrps16 intron phylogeny of the tribeSileneae (Caryophyllaceae)

Intron sequences of the chloroplast generps16 from 46 species were used to examine phylogenetic relationships indicated by nrDNA ITS sequence variation in the tribeSileneae (Caryophyllaceae, Caryophylloideae). This region has previously not been utilized for phylogenetic purposes but the results presented here suggest that it is a consistent and valuable complement to the ITS sequences. Therps16 intron trees are largely congruent with the ITS trees. All the major hypotheses suggested by the ITS data are supported, often at similar bootstrap levels. The joint usage ofrps16 intron and ITS sequences provides a powerful tool for resolving many of the difficult taxonomic issues in the tribeSileneae. Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Thermoregulation in three species of Afrotropical subterranean mole-rats (Rodentia: Bathyergidae) from Zambia and Angola and scaling within the genus Cryptomys

The thermoregulatory characteristics of three species of Cryptomys from Zambia and Angola are examined and, together with published data on four other species of Cryptomys from southern Africa, used to determine whether scaling occurs in this genus of subterranean rodents. The thermoregulatory properties of acclimated giant Zambian mole-rats, Cryptomys mechowi ( =267 g), Angolan mole-rats, Cryptomys bocagei ( =94 g) and Zambian common mole-rats Cryptomys hottentotus amatus ( =77 g) are as follows. Mean resting metabolic rates (RMRs) within the respective thermoneutral zones were 0.60±0.08 cm³ O₂ g^-1 h^-1 (n=12) for C. mechowi; 0.74±0.06 cm³ O₂ g^-1 h^-1 (n=8) for C. bocagei and 0.63±0.06 cm³O₂ g^-1 h^-1 (n=21) for C. h. amatus. The thermoneutral zones (TNZs) of all three species are narrow: 29–30°C for C. mechowi; 31.5–32.5°C for C. bocagei and 28–32° C for C. h. amatus. The increase in mean RMR at the lowest temperatures tested (15° C for C. mechowi, 18° C for C. bocagei and C. h. amatus) was 2.35, 2.2 and 3.82 times their RMR in the TNZ respectively. Body temperatures are low, 34±0.53° C (n=24) for C. mechowi, 33.7±0.32° C (n=20) for C. bocagei and 33.8±0.43° C (n=40) for C. h amatus. At the lower limit of thermoneutrality, conductances are 0.09±0.01 cm³ O₂ g^-1 h^-1 °C^-1 (n=30) in C. mechowi; 0.12±0.01 cm³ O₂ g^-1 h^-1 °C^-1 (n=20) in C. bocagei and 0.12±0.03 cm³ O₂ g^-1 h^-1 °C^-1 (n=32) in C. h. amatus. The range in mean body mass among the seven species of Cryptomys examined for scaling was 60 g (C. darlingi) to 267 g (C. mechowi). There is no clear relationship between RMR within the TNZ and body mass. The resultant relationship is represented by the power curve RMR=2.45 mass^-0.259.     

A test of a mitochondrial gene-based phylogeny of woodpeckers (genus Picoides) using an independent nuclear gene, beta-fibrinogen intron 7

A conservative estimate of the species tree for the woodpecker genus Picoides based on two mitochondrial protein-coding genes is tested using sequences of an independently evolving nuclear intron, beta-fibrinogen intron 7. The mitochondrial gene-based topology and the intron-based topology are concordant, and a partition-homogeneity statistical test did not detect phylogenetic heterogeneity. The intron evolves more slowly than the mitochondrial sequences and tends not to resolve relationships among recently evolved species. However, the intron is superior over mitochondrial genes in resolving older bifurcations in the phylogeny. The two data sets were combined resulting in a robust estimate of the Picoides species tree in which most every node is statistically supported by bootstrap proportions. The Picoides species tree clearly shows that many morphological and behavioral characters used to lump species into this single genus have evolved by convergent evolution. Picoides is considered the largest genus of woodpeckers, but the molecular-based species tree suggests that Picoides is actually a conglomerate of several smaller groups.     

Therps16 intron and the phylogeny of theRubioideae (Rubiaceae)

The phylogeny of the subfamilyRubioideae (Rubiaceae) was estimated from sequence variation in therps16 intron (cpDNA) in 143 ingroup and 5 outgroup taxa. The analysis largely confirms a recent one based onrbcL sequences, but branch support is often much stronger. Three of the traditional subfamilies are supported,Rubioideae, Cinchonoideae s. str., andIxoroideae s. l. while there is no support forAntirheoideae. TheRubioideae are the sister group of all otherRubiaceae and comprise the tribesAnthospermeae, Coccocypseleae, Cruckshanksieae, Coussareeae, Gaertnereae, Hedyotideae, Knoxieae, Morindeae, Ophiorrhizeae, Paederieae, Pauridiantheae, Perameae, Psychotrieae, Rubieae, Spermacoceae, Theligoneae, andUrophylleae. TheHamelieae andHillieae belong to theCinchonoideae. Rachicallis andSiemensia should be transferred from theHedyotideae to theCinchonoideae. ThePauridiantheae, Urophylleae, Ophiorrhizeae, andRaritebe form the basalmost subclade of theRubioideae. The second basalmost clade consists of the generaLasianthus andPerama. The third basalmost clade consists of the tribesCoussareeae, Coccocypseleae andCruckshanksieae, and the generaDeclieuxia andHindsia. The tribesKnoxieae, Anthospermeae, Argostemmateae, Paederieae, Theligoneae, Rubieae, Hedyotideae, andSpermacoceae are members of one clade. TheKnoxieae are monophyletic ifOtiophora, Otomeria, andPentas are included. The tribeAnthospermeae is supported as monophyletic, but its subtribes are not. ThePaederieae, together withTheligonum, form a paraphyletic grade basal to theRubieae. TheHedyotideae, includingSchismatoclada, form a grade at the base of theSpermacoceae. TheGaertnereae are monophyletic and distinct from thePsychotrieae. TheMorindeae are monophyletic and includeDamnacanthus andMitchella. Schradera is the sister group of theMorindeae. ThePsychotrieae are monophyletic when theGaertnereae, Lasianthus, andDeclieuxia are excluded. The recognition of a subtribeHydnophytineae leaves the rest of thePsychotrieae paraphyletic.Psychotria is paraphyletic with respect to all other genera of the tribe. Approximately 50 genera are here classified for the first time based on molecular data.     

Molecular phylogeny of Fordini (Hemiptera: Aphididae: Pemphiginae) inferred from nuclear gene EF-1 alpha and mitochondrial gene COI

The tribe Fordini is a fascinating group because of its complicated life history, primary host specificity and gall-forming characteristic. Different species produce galls with different morphology on different parts of the host plants. The EF-1alpha-based, COI-based and combined sequences-based phylogenetic trees with three algorithms MP, ML and Bayes all strongly suggest that Fordini is a monophyletic group with two clades corresponding to two subtribes, Fordina and Melaphidina, each also monophyletic. Some important morphological characters and primary host plants of aphids were mapped onto the phylogenetic tree to analyse the division of subtribes and to uncover at which level the aphids correspond to their primary hosts, Pistacia and Rhus. Results suggest that the division of subtribes in Fordini is closely related to host selection of aphids. The evolution of gall morphology and the probable driving force behind it in this tribe were also discussed. The Fordini aphids seem to have evolved towards a better ability to manipulate their host plant, induce strong sinks and gain high reproductive success. Galls in this tribe evolved mainly along two directions to attain this goal: (i) by enlarging the gall from small bag to spherical, even big cauliflower-like, and changing the galls' location or forming two galls in their life cycle (Fordina); (ii) by moving the gall position from midrib, petiole of the leaflet, and eventually to the common petiole of the compound leaf (Melaphidina).