Social wasps,crickets and cockroaches contribute to pollination of the holoparasitic plant Mitrastemon yamamotoi (Mitrastemonaceae) in southern Japan

• Mitrastemon yamamotoi is completely embedded within the tissues of its hosts, except during the reproductive stage, when aboveground parts emerge from host tissues. Its highly modified appearance has attracted attention of many botanists, but very little is known about the reproductive system.
• Floral visitors to M. yamamotoi were observed in southern Japan. Pollination experiments were conducted to determine the plant's self‐compatibility and pollen limitation, as well as the contribution of diurnal and nocturnal visitors to fruit set and outcrossing.
• Mitrastemon yamamotoi is mainly pollinated by social wasps, but previously unnoticed pollinators (i.e. crickets and cockroaches) are also important, based on visitation frequency and pollen loads. Results of the pollination experiments suggest that nocturnal visitors, such as crickets and cockroaches, contribute to geitonogamous pollination, whereas diurnal visitors, such as social wasps, facilitate outcrossing.
• The unexpected pollinator assemblage of M. yamamotoi might be influenced by multiple factors, including the highly modified flowers that are produced close to the ground in dark understorey environments, the species’ winter‐flowering habit and the location of the study site (i.e. near the northern limit of the species’ range). Considering that M. yamamotoi occurs widely in subtropical and tropical forests in Asia, additional studies are needed to assess pollinator assemblages of M. yamamotoi at other locations.

     

Fruit‐ and seed‐feeding habit of the crane fly Libnotes puella (Diptera: Limoniidae) in Ryukyu Islands,Japan

Although most crane fly larvae consume decaying plant materials and their associated microorganisms, all Libnotes species investigated so far are known to be xylophilous insects. Here we report the novel herbivorous feeding ecology of Libnotes puella. We found that L. puella larvae consumed the fruits or seeds of three unrelated plants, Mitrastemon yamamotoi (Mitrastemonaceae), Balanophora tobiracola (Balanophoraceae) and Barringtonia racemosa (Lecythidaceae). The larvae live within dense tubes formed by their feces fastened by their mucous excretions at the final stage, and they pupate within such tubes. This is the first detailed documentation of the fruit‐ and seed‐feeding habit of crane flies.     

Simulated Intraspecific Usurpations in Paper Wasps: Different Reproductive Tactics Affect Foreign Brood Destruction in Polistes fuscatus and Polistes dominulus

We simulated intraspecific usurpation in two species of paper wasps at a field site in Michigan to compare the species' treatment of foreign brood. Queens of Polistes fuscatus, a species that commonly uses intraspecific usurpation as an alternative reproductive tactic, destroyed significantly fewer large larvae and significantly more small larvae than queens of P. dominulus, a sympatric species that rarely usurps. The pattern of brood destruction exhibited by P. fuscatus was consistent with the previously published findings that P. fuscatus usurpers destroy reproductive‐destined brood (eggs and small larvae), but not worker‐destined brood (large larvae and pupae) that are subsequently used by a usurper to raise her own reproductives. The pattern of brood destruction displayed by P. dominulus differed from that of Polistes species that frequently engage in intraspecific usurpation. The brood destruction pattern in P. dominulus may have been shaped by nest adoption, a common alternative reproductive tactic in this species. If so, it is not clear why P. dominulus would destroy large, worker‐destined larvae.     

Gregarious emergence of Macrostomion sumatranum (Hymenoptera: Braconidae; Rogadinae) from the mummified, full-grown larvae of Theretra silhetensis (Lepidoptera: Sphingidae)

Twenty‐six to 160 adults of Macrostomion sumatranum emerged from a mummified full‐grown larva of a sphingid moth, Theretra silhetensis, on taro in Okinawa‐jima, Japan. This is the first host record of M. sumatranum. The sex ratio was female‐biased in a small brood but less so or even male‐biased in a large brood.     

The ontogeny of handling hard‐to‐process food in wild brown capuchins (Cebus apella apella): evidence from foraging on the fruit of Maximiliana maripa

We examined age‐related differences in wild brown capuchins' foraging efficiency and the food‐processing behaviors directed toward maripa palm fruit (Maximiliana maripa). A detailed comparison of the different foraging techniques showed that plucking the fruit from the infructescence constituted the main difficulty of this task. Foraging efficiency tended to increase with age, with a threshold at which sufficient strength allowed immatures by the age of three to reach adult‐level efficiency. Youngsters spent more time than older individuals browsing the infructescence and pulling the fruit in an attempt to harvest it. Infants tried to compensate for their inability to pluck fruit by adopting alternative strategies but with low payback, such as gnawing unplucked fruit and opportunistically scrounging others' partially processed food. Although around 2 years of age, young capuchins exhibited all of the behaviors used by adults, they did not reach adult‐level proficiency at feeding on maripa until about 3 years (older juveniles). We compared this developmental pattern with that of extractive foraging on beetle larvae (Myelobia sp.) hidden in bamboo stalks, a more difficult food for these monkeys [Gunst N, Boinski S, Fragaszy DM. Behaviour 145:195–229, 2008]. For maripa, the challenge was mainly physical (plucking the fruit) once a tree was encountered, whereas for larvae, the challenge was primarily perceptual (locating the hidden larvae). For both foods, capuchins practice for years before achieving adult‐level foraging competence, and the timeline is extended for larvae foraging (until 6 years) compared with maripa (3 years). The differing combinations of opportunities and challenges for learning to forage on these different foods illustrate how young generalist foragers (i.e. exploiting a large number of animal and plant species) may compensate for their low efficiency in extractive foraging tasks by showing earlier competence in processing less difficult but nutritious foods, such as maripa fruit. Am. J. Primatol. 72:960–973, 2010. © 2010 Wiley‐Liss, Inc.     

Bionomics and rearing of Miltotranes prosternalis (Lea, 1929) (Coleoptera: Curculionidae), a mutualistic cycad pollinator in Australia

Pollination is the crucial stage for reproduction and survival of plants, therefore knowledge about the bionomics of pollinators is often critical for plant conservation. Bowenia spectabilis (Cycadales: Stangeriaceae) is a CITES-protected cycad species endemic to the Wet Tropics Bioregion of Queensland, Australia. It is pollinated by a host-specific weevil species, Miltotranes prosternalis (Lea, 1929) (Curculionidae: Molytinae), which breeds in its male cones but also visits its female cones and pollinates them. The association between the cycad and the weevil is an obligate brood-site reward pollination system, in which the weevil is provided with food and a brood site by the plant in “return” for the service of pollinating it. To facilitate the knowledge of the natural history of this unique Bowenia pollinator, we developed a method for rearing mature larvae of M. prosternalis collected from the field in the laboratory. Furthermore, we summarize biological information of M. prosternalis gathered from previous studies and by firsthand observations in the wild and during rearing in captivity.     

First fossil record of an East Asian endemic genus Sladenia (Sladeniaceae) from its modern range: Implications for floristic evolution and conservation biology

Fossil records of endemic plants play an important role in recognizing the floristic history of East Asia and thereby facilitate the conservation of plant diversity in the region. However, the fossil record of many extant East Asian endemic genera remains poorly documented thus far. Here, we report an infructescence fossil of an East Asian endemic genus, Sladenia (Sladeniaceae), from the early Miocene of southeastern Yunnan, China. The fossil is characterized by: (i) dichasial cymes; and (ii) flask‐shaped ovary with dense subparallel ribs on the surface extending from the base to the distal end of the united style. It represents the first fossil record of Sladenia in Asia, showing that the genus was established in the region at least by the early Miocene. Given that a much older fossil record of Sladeniaceae has been reported from Africa and the sister group of Sladenia is distributed only in Africa, Sladenia is not likely of East Asian origin. The present endemic status of Sladenia was possibly achieved by regional extirpation in Africa and taking refuge in East Asia. This case thus supports the “Museum” rather than “Cradle” hypothesis for the genesis of high plant species in the flora of East Asia. A comparison of the present fossil with extant Sladenia infructescence shows morphological stasis from the early Miocene to present. Such evolutionary tardiness might have resulted in the reduced fitness of the genus, which further caused its current endangered situation.     

Mating behavior,spawning, parental care,and embryonic development of some marine pseudocerotid flatworms (Platyhelminthes: Rhabditophora: Polycladida) in Singapore

The reproductive biology of tropical marine cotylean polyclads is presently poorly known. Reproductive aspects of 16 polyclad species from the family Pseudocerotidae in the genera Acanthozoon, Nymphozoon, Phrikoceros, Pseudobiceros, Pseudoceros, Thysanozoon, and Tytthosoceros from Singapore were documented for the first time. Penis fencing was observed to be just a mating ritual and not necessary for insemination, not always aggressive, and could also result in eventual reciprocal insemination. Results showed that all species underwent similar embryonic developmental stages and hatched as Müller's larvae. Only Pseudoceros concinnus and Pseudoceros laingensis, with mean developmental times of >20 days and mean brood sizes of <1,000 eggs, displayed long‐term parental care. Polyclads producing larger brood sizes had shorter developmental times and only covered their egg masses for about one day. Phrikoceros baibaiye and Pseudobiceros spp. produced egg capsules with pointed opercula, whereas all other species possessed smooth, rounded opercula. All genera hatched with brownish‐orange larvae, except for Pseudoceros spp., which hatched with reddish‐purple larvae regardless of the initial embryo color (either grayish‐yellow or reddish‐purple). These could potentially complement current taxonomic characters in distinguishing polyclad genera and species.     

Host defense in Nicrophorus quadripunctatus against brood parasitism by Ptomascopus morio (Coleoptera: Silphidae: Nicrophorinae)

Few studies have been conducted on the host defenses of insects against brood parasitism. We investigated whether the silphid beetle Ptomascopus morio, a brood parasite of related silphid species Nicrophorus concolor, can also parasitize another silphid species Nicrophorus quadripunctatus and the manner in which N. quadripunctatus defends itself against parasitism. Successful brood parasitism under natural conditions was not observed at the time of year when P. morio and N. quadripunctatus are both reproductively active. Follow-up experiments revealed that P. morio attempts to oviposit near N. quadripunctatus nests, but is rarely successful if adult hosts are present. When P. morio larvae were experimentally introduced to N. quadripunctatus broods, some P. morio larvae survived when the host and parasite larvae were at the same stage. We concluded that N. quadripunctatus defends itself against brood parasitism in two ways: (1) potential brood parasites are repelled, thus limiting their access to the resource; and (2) the young of the parasitic species are killed.     

Root damage to apple plants by cockchafer larvae induces a change in volatile signals below‐ and above‐ground

Volatile organic compounds (VOCs) mediate communication between plants and insects. Plants under insect herbivore attack release VOCs either at the site of attack or systemically, indicating within‐plant communication. Some of these VOCs, which may be induced only upon herbivore attack, recruit parasitoids and predatory insects to feed on the attacking insects. Moreover, some plants are able to ‘eavesdrop’ on herbivore‐induced plant volatiles (HIPVs) to prime themselves against impending attack; such eavesdropping exemplifies plant–plant communication. In apple orchards, the beetle Melolontha melolontha L. (Coleoptera: Scarabaeidae) is an important insect pest whose larvae live and feed on roots for about 4 years. In this study, we investigated whether the feeding activity of M. melolontha larvae (1) alters the volatile profile of apple roots, (2) induces the release of HIPVs systemically in the leaves, and (3) whether infested plants communicate to neighbouring non‐infested conspecifics through HIPVs. To answer these questions, we collected constitutive VOCs from intact M9 roots as well as M. melolontha larvae‐damaged roots using a newly designed ‘rhizobox’, to collect root‐released volatiles in situ, without damaging the plant root system. We also collected VOCs from the leaf‐bearing shoots of M9 whose roots were under attack by M. melolontha larvae and from shoots of neighbouring non‐infested conspecifics. Gas chromatography‐mass spectrometry analysis showed that feeding activity of M. melolontha larvae induces the release of specific HIPVs; for instance, camphor was found in the roots only after larvae caused root damage. Melolontha melolontha also induced the systemic release of methyl salicylate and (E,E)‐α‐farnesene from the leaf‐bearing shoots. Methyl salicylate and (E,E)‐α‐farnesene were also released by the shoots of non‐infested neighbouring conspecifics. These phenomena indicate the induction of specific VOCs below‐ and above‐ground upon M. melolontha larvae feeding on apple roots as well as plant–plant communication in apple plants.     

Oviposition and mining by Phytobia betulae (Diptera: Agromyzidae) in genotypes of European white birch (Betula pendula)

Abstract 1 The colonization success of herbivorous insects depends partly on the ability of females to choose suitable host plants. Phytobia betulae Kangas (Diptera: Agromyzidae), a miner of differentiating xylem, infests birch (Betula pendula Roth and Betula pubescens Ehrh.) trees. 2 In a field experiment we measured successful ovipositions in the long shoots of eight 5‐year‐old‐birch genotypes. Later in the growing season, in the same experiment, we counted the number of larvae from the same birch genotypes (different replicates). 3 We attempted to determine where Phytobia females lay their eggs in the birch canopy, what kind of shoots Phytobia utilize, whether birch genotypes differ in their susceptibility to Phytobia, and which tree characteristics correlate with successful utilization of birch for oviposition and larval development. 4 The ovipositions were concentrated in fast growing shoots and only 5% of the available shoots were utilized. Usually a shoot contained only one oviposition (80% of the cases). However, oviposition pattern resulted in significant aggregations (two to four ovipositions per shoot): 11% of shoots selected for oviposition contained more than one Phytobia and 22% of larvae shared their shoot with at least one other larva. Although birch shoots were much more abundant than Phytobia, there may be intraspecific competition for host resources among Phytobia. 5 Birch genotypes differed in the number of ovipositions but not in the number of larvae, although in general the number of larvae was correlated positively with the number of ovipositions. There were more ovipositions in the faster growing genotypes.     

Determining the host‐plant resistance mechanisms for Mamestra brassicae (Lepidoptera: Noctuidae) pest in cabbage

Six cabbage (Brassica oleracea var. capitata) varieties with different levels of resistance to Mamestra brassicae (Lepidoptera: Noctuidae) were investigated in order to assess whether antibiosis and antixenosis mechanisms are involved in the resistance to this pest or not. Four experiments were conducted to determine the effect of variety and plant ontogeny on larval behaviour, adult oviposition and leaf damages in non‐choice and choice tests. Larval survival, time to development and larval weights differed depending on the varieties and plant stages that we tested. At the pre‐head stage, larval mortality was higher, larvae died faster, time to pupation was shorter, pupae were lighter and the percentage of viable pupae and growth index (GI) values were lower than larvae reared from plants at the head stage. The commercial hybrid ‘Corazón de buey’ and the local variety named ‘BRS0535’ exhibited antibiosis to M. brassicae as they reduced its survival and growth and delayed its development time. In addition, these varieties were the most resistant after artificial infestation in terms of head foliage consumption and number of larvae per plant. Oviposition tests demonstrated that resistance found in ‘Corazón de buey’ and BRS0535 could be also based on antixenosis mechanisms as they resulted in fewer egg batches on plants, whereas BRS0402 could be classified as resistant because M. brassicae larvae showed less preference for it. Thus, resistance to M. brassicae found in cabbage crops may be due to the joint action of several factors involving antibiosis and antixenosis. We found significant differences in the resistance of BRS0535 depending on the plant ontogeny as it loses its resistance while developing. Further studies are required to identify the mechanism of antibiotic resistance which is present in this variety at the pre‐head stage and the changes that occur in plant defence as it grows.     

Oviposition and larval dispersal of the common armyworm,Mythimna convecta (Walker) (Lepidoptera: Noctuidae)

The common armyworm, Mythimna convecta (Walker), is a pest of cereal crops and pasture grasses in Australia. During autumn, egglaying in grasslands commonly occurs before plant growth commences. The possible association between oviposition and dried grasses was investigated in field studies of larval distribution in a pasture and a crop habitat, and in laboratory studies of oviposition site preferences and the mode of dispersal of newly hatched first instar larvae. A comparison of a green grass pasture with and without a component of dried grass showed that highest densities of M. convecta larvea. were found in the former. In the laboratory, egg batches were laid between plant parts in close contact, such as between the stem and leaf axil and between seeds in the seed head. The crevices in dried plant material and seed-heads were significantly more acceptable for oviposition than in green foliage. Oviposition in dried plant material enables M. convecta to rapidly colonize ephemeral grasslands immediately after rains when the neonate larvae can disperse and feed on new vegetative growth. Shortly after hatching, 93% of first instar larvae used fine silken threads for aerial dispersal in 0.5 m s^-1 air currents and 43% were blown a distance of more than 1.0 m. In the field, sticky traps caught larvae dispersing in all directions from a central source, and indicated that approximately 30% were transported 2 m after 7 days of light to medium breezes. Terminal velocities of newly hatched larvae were 20–130 cm s^-1, depending on the length of the trailing thread.     

Host‐plant use by two Orthomeria (Phasmida: Aschiphasmatini) species feeding on Macaranga myrmecophytes

Myrmecophytes depend on symbiotic ants (plant‐ants) to defend against herbivores. Although these defensive mechanisms are highly effective, some herbivorous insects can use myrmecophytes as their host‐plants. The feeding habits of these phytophages on myrmecophytes and the impacts of the plant‐ants on their feeding behavior have been poorly studied. We examined two phasmid species, Orthomeria alexis and O. cuprinus, which are known to feed on Macaranga (Euphorbiaceae) myrmecophytes in a Bornean primary forest. Our observations revealed that: (i) each phasmid species relied on two closely‐related myrmecophytic Macaranga species for its host‐plants in spite of their normal plant‐ant symbioses; and (ii) there was little overlap between their host‐plant preferences. More O. cuprinus adults and nymphs were found on new leaves, which were attended by more plant‐ants than mature leaves, while most adults and nymphs of O. alexis tended to avoid new leaves. In a feeding choice experiment under ant‐excluded conditions, O. alexis adults chose a non‐host Macaranga myrmecophyte that was more intensively defended by plant‐ants and was more palatable than their usual host‐plants almost as frequently as their usual host‐plant, suggesting that the host‐plant range of O. alexis was restricted by the presence of plant‐ants on non‐host‐plants. Phasmid behavior that appeared to minimize plant‐ant attacks is described.     

Oogenesis and larval development in Micromaldane spp. (Polychaeta: Capitellida: Maldanidae)

Summary

The morphology of larval development and some aspects of oogenesis are described in Micromaldane spp. from eastern Australia. All four species breed throughout the year, and in all the oocytes are released into the coelom prior to vitellogenesis. Micromaldane androgyne, M. pamelae and M. rubrospermatheca all have solitary oogenesis, while Micromaldane nutricula has “nurse” cells attached to the oocytes throughout vitellogenesis. Mature females of Micromaldane nutricula brood up to two groups of 1–4 directly developing larvae at a time. Newly fertilized eggs are ellipsoidal, 150x350 μm. No ciliation is visible during development. Larvae leave the tube after developing 12 chaetigers. All Micromaldane androgyne adults brood a single batch of two or three directly developing larvae at a time. Newly deposited eggs are 1 mm long and 150 μm in diameter. No ciliation is visible on the larvae during development. Larvae develop 19 chaetigers before leaving the tube. Mature females of Micromaldane pamelae brood single batches of up to 31 larvae at a time. Mature oocytes are ellipsoidal 300 X 150 μm. Larvae show a distinct neurotroch in early stages of development, but this is resorbed as chaetigers are formed. Larvae probably leave the tube at the 16-chaetiger stage.

Comparisons are made with other polychaetes and members of the Capitellida. Micromaldane spp. may be useful in the study of reproductive effort and covariables, outlined for polychaetes by Olive (1985), such as body size with longevity and fecundity (inversely) with brood frequency.     

Direct and indirect effects of predatory wasps (Polistes sp.: Vespidae) on gregarious caterpillars (Hemileuca lucina: Saturniidae)

Summary We examined how predation by vespid wasps,Polistes dominulus andP. fuscatus, affected the behavior, growth rate and survivorship of aggregated caterpillars ofHemileuca lucina (Saturniidae). Although these larvae can exhibit a variety of defense and escape behaviors, in general larvae reacted to wasp attacks by clinging to the hostplant. Neighboring larvae in the aggregation responded by leaving the feeding site and moving to the interior or base of the plant. To determine wheter wasp attack affected the behavior and growth of the caterpillars that escaped, a field experiment was conducted with treatments of: 1) larvae exposed to wasps, 2) larvae protected from wasps, and 3) larvae protected from wasps but with the attack of wasps simulated (=harassment). Over just one instar, protected larvae gained significantly more weight than the harassed larvae, which in turn weighed significantly more than the larvae that escaped the wasps. The behavior of attacked and harassed larvae differed from that of the protected larvae; the disturbed larvae often fed in smaller groups and in shaded portions of the plant where only mature leaves were available. A laboratory experiment showed that at 35° C (daytime temperature) larvae had significantly higher relative growth rates and significantly shorter instar duration than larvae reared at 25° C. Our results suggest that wasps, in addition to killing caterpillars, indirectly affect larval fitness by slowing larval growth, at least in part by forcing larvae into cooler microhabitats where leaves are of lower quality.     

Suitability of Eucalyptus and Corymbia for Mnesampela privata (Guenée) (Lepidoptera: Geometridae) larvae

Abstract 1 Mnesampela privata (Guenée) has a host list of 40 Eucalyptus and at least one Corymbia species. Larval survival and performance was studied on 19 species to investigate how certain leaf traits influence the suitability of different species. 2 After 7 days, survival on Eucalyptus aggregata and Eucalyptus camphora is greater than 70% even though the toughness of leaves is 0.15–0.19 mg/mm². However, after the same time, survival on genotypes of Eucalyptus melliodora and Eucalyptus sideroxylon was less than 60%, or even 0%, even though the toughness of some leaves was as low as 0.11 mg/mm². An unmeasured allelochemical, rather than toughness, may reduce survival on these species. 3 Dry weights of first‐instar larvae were negatively correlated with leaf toughness for 13 of the species studied. Species that produced the heaviest first‐instar larvae were not the same hosts that produced the heaviest second‐instar larvae. 4 Dry weights of female pupae were negatively correlated with total oil content for five of the species studied. 5 Larvae exhibit age‐related changes in feeding behaviour. Neonates skeletonize leaves (avoid leaf veins and oil glands) and post‐third‐instar larvae ingest whole leaf fragments (consume small leaf veins and oil glands). These findings suggest that neonates are sensitive to high leaf toughness and non‐oil plant secondary metabolites whereas older larvae are less sensitive to high leaf toughness and are likely to become larger adults on hosts with lower oil contents.     

Reproductive trade-offs in a specialized plant/pollinator system involving Asplundia uncinata Harling (Cyclanthaceae) and a derelomine flower weevil (Coleoptera: Curculionidae)

The reproductive interactions of a specialized plant/pollinator system involving Asplundia uncinata Harling (Cyclanthaceae) and a derelomine flower weevil (Coleoptera: Curculionidae: Derelomini) were studied at La Selva, Costa Rica. The inflorescences of A. uncinata exhibit a suite of cantharophilous characters including a precisely synchronized anthesis, protogyny, thermogenesis, olfactory attraction of visitors via modified staminodes, and narrow interfloral entrances. The weevil pollinators use the inflorescences for feeding, mating, and oviposition. The larvae are detritivorous and develop either in the detaching staminate flowers or, at a more favorable rate, in the rotting infructescences. The rate of infructescence abortion was high and caused by low levels of pollination. Manual pollination treatments yielded significantly higher seed counts than obtained under natural conditions, and furthermore demonstrated the inflorescences' ability to reproduce via geitonogamy. In the longer term, the reproductive benefits of maintaining low levels of pollination may shift away from the weevils and towards the plants via an increase in the size of the pollinator population.     

Rate heterogeneity in six protein-coding genes from the holoparasite Balanophora (Balanophoraceae) and other taxa of Santalales

Background and Aims

The holoparasitic flowering plant Balanophora displays extreme floral reduction and was previously found to have enormous rate acceleration in the nuclear 18S rDNA region. So far, it remains unclear whether non-ribosomal, protein-coding genes of Balanophora also evolve in an accelerated fashion and whether the genes with high substitution rates retain their functionality. To tackle these issues, six different genes were sequenced from two Balanophora species and their rate variation and expression patterns were examined.

Methods

Sequences including nuclear PI, euAP3, TM6, LFY and RPB2 and mitochondrial matR were determined from two Balanophora spp. and compared with selected hemiparasitic species of Santalales and autotrophic core eudicots. Gene expression was detected for the six protein-coding genes and the expression patterns of the three B-class genes (PI, AP3 and TM6) were further examined across different organs of B. laxiflora using RT-PCR.

Key Results

Balanophora mitochondrial matR is highly accelerated in both nonsynonymous (d_N) and synonymous (d_S) substitution rates, whereas the rate variation of nuclear genes LFY, PI, euAP3, TM6 and RPB2 are less dramatic. Significant d_S increases were detected in Balanophora PI, TM6, RPB2 and d_N accelerations in euAP3. All of the protein-coding genes are expressed in inflorescences, indicative of their functionality. PI is restrictively expressed in tepals, synandria and floral bracts, whereas AP3 and TM6 are widely expressed in both male and female inflorescences.

Conclusions

Despite the observation that rates of sequence evolution are generally higher in Balanophora than in hemiparasitic species of Santalales and autotrophic core eudicots, the five nuclear protein-coding genes are functional and are evolving at a much slower rate than 18S rDNA. The mechanism or mechanisms responsible for rapid sequence evolution and concomitant rate acceleration for 18S rDNA and matR are currently not well understood and require further study in Balanophora and other holoparasites.