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Two new species of Schistonchus Tylenchida Aphelenchoididae associated with Ficus macrophylla from Australia. Fundamental and Applied Nematology, 201: 79-86
Abstract
Two new species of the genus Schistonchus are described from inflorescences of Ficus macrophylla in Australia: S. macrophylla n. sp. from Adelaide and S. altermacrophylla n. sp. from Sydney. S. macrophylla n. sp. is characterized by the tail without a mucron and the excretory pore at the level of the median bulb in both sexes, spicule shape in the males, and a long post-vulval sac in females. S. altermacrophylla n. sp. is characterized by having the excretory pore very close to head region. The diagnosis of the genus is emended. Both new species develop concomitantly with the wasp Pleistodontes froggatti. S. macrophylla n. sp. was carried between inflorescences in the abdomen of the wasp. S. macrophylla n. sp. was also recovered from three other species of Ficus, all endemic to Australia.
... Cobb, 1927. So far, 27 species are classified in the genus Ficophagus (Lloyd & Davies, 1997;DeCrappeo & Giblin-Davis, 2001;Zeng et al., 2007Zeng et al., , 2010Zeng et al., , 2011Zeng et al., , 2013Zeng et al., , 2019Zeng et al., , 2020Bartholomaeus et al., 2009Bartholomaeus et al., , 2012Davies et al., 2010Davies et al., , 2013Davies et al., , 2015Davies et al., , 2017aDavies et al., , b, 2020Bajaj & Tomar, 2014;Zhao et al., 2015;Zhang et al., 2019;Gupta et al., 2021;Zhao et al., 2022). Recent survey work on the diversity of fig nematodes in China revealed an undescribed species of nematode from F. annulata in Xishuangbanna, Yunnan Province, P.R. China. ...
... By having similarity in PUS (0.4 VBD long), the new species was compared with 18 known species, namely: F. aculeate; F. altermacrophylla (Lloyd & Davies, 1997 (Bartholomaeus, Davies, Ye, Kanzaki & Giblin-Davis, 2009) Davies & Bartholomaeus, 2015;and F. yoponensis Davies, Ye, Center, Kanzaki, Bartholomaeus, Herre, Esquivel & Giblin-Davis, 2017. ...
... The new species differs as follows: from F. aculeata by possessing a different relative EP position (at level between stylet basal knobs and metacorpus vs near the lips), male tail tip morphology (without mucron vs with), a longer body (L = 530-696 vs 396-523 μm in males, 755-847 vs 548-671 μm in reproductive females), a larger spicule (28-29 vs 15-17 μm), a lower ratio c (1.5-1.8 vs 1.8-2.8) in males, a higher ratio c (3-4 vs 2.0-3.1) in reproductive females (Davies et al., 2010); from F. altermacrophylla by possessing a different relative EP location (at level between stylet basal knobs and metacorpus vs just posterior to the head), female tail tip (with mucron vs without), male tail tip morphology (without mucron vs with), papilla arrangement (one pair just adcloacal, one pair halfway between cloacal aperture and tail terminus and one pair near tail tip vs one pair near rostrum, one pair adcloacal, and one pair halfway between cloacal aperture and tail terminus), a longer body (L = 755-847 vs 411-571 μm) in reproductive females, a larger c value (18-23 vs 13-16) (Lloyd & Davies, 1997); from F. altissimus by possessing a different relative EP position (at level between stylet basal knobs and metacorpus vs near the lips), reproductive female tail tip morphology (without mucron vs with), spicule morphology and size (bow-shaped without cucullus vs hook-shaped with, 19-20 vs 12-16 μm), and a longer male body (L = 755-847 vs 424-515 μm) (Zeng et al., 2013); from F. aureus by possessing a different relative EP position (at level between stylet basal knobs and metacorpus vs near the head), spicule morphology ( (Davies et al., 2017b); from F. chaozhouensis by possessing a different relative EP position (at level between stylet basal knobs and metacorpus vs near the head), spicule morphology (bow-shaped, rostrum indistinct vs hammer head-shaped, rostrum distinct, stout with broadly squared tip), male tail tip morphology (without mucron vs with), a smaller V value (71-73 vs 74.5-78.5), a higher ratio a (23-26 vs 13.9-19.0 in male; 23-29 vs 16.2-22.6 in female), a larger spicule (28-29 vs 22-24 μm), and a longer reproductive female body (L = 755-847 vs 510-662 μm) (Zeng W. et al., 2020); from F. costaricanus by possessing a different relative EP position (at level between stylet basal knobs and metacorpus vs near the head), a larger spicule (28-29 vs 13-17 μm) (Davies et al., 2017b); from F. giblindavisi by possessing a different relative EP position (at level between stylet basal knobs and metacorpus vs at or posterior to the nerve ring), tail tip (without mucron vs with), spicule morphology (indistinct rostrum vs distinct; without cucullus vs with), a shorter stylet (17-18 vs 35.1-45.8 μm in males, 17-18 vs 29.2-34.2 ...
A new species of the genus Ficophagus was recovered from the syconia of Ficus annulata from Xishuangbanna, Yunnan province, China. It is described herein as Ficophagus annulatae n. sp. and is characterised by possessing the combined characters of a short PUS (9-18 μ m or 0.3-0.6 VBD long), an excretory pore (EP) located at the level between stylet basal knobs and metacorpus, presence of crustaformeria, amoeboid sperm, three pairs of subventral papillae on the male tail (one pair just adcloacal (P2), one pair halfway between cloacal aperture and tail terminus (P3), and one pair near tail tip (P4)), rounded male tail tip without mucron, absence of gubernaculum and bow-shaped spicule with indistinct rostrum. Ficophagus annulatae n. sp. was differentiated from other sequenced species by the partial small subunit (SSU) rRNA gene and D2-D3 expansion segments of the large subunit (LSU) rRNA gene. Phylogenetic analysis with the LSU D2-D3 expansion segment sequence suggested that F. annulatae n. sp. is clustered in the same highly supported monophyletic clade with F. benjamina , F. curtipes and F. microcarpus . It differs morphologically from these species in EP position, spicule and uterus morphology, and some morphometric characters.
... Nematodes were collected from sycones as described in Lloyd & Davies (1997). Latitude and longitude were determined using a Magellan ® GPS 315/320 or GPS Location COP-APPS.net. ...
... Based on SSU molecular studies (Fig. 1), Schistonchus macrophylla Lloyd & Davies, 1997 appears to form a species complex (GenBank number KY996313), sister to, but different from, S. superbus Zeng, Ye, Li, Wang, Du & Giblin-Davis, 2013b (HM151003, 24 bp differences in 1510 bp), S. caprifici Gasparrini, 1874 (FN564938, 18 Although the Bayesian analysis of the D2-D3 fragment of the LSU sequences involved a different subset of nematodes to the SSU analysis (Fig. 2), the results are generally similar to those using the more conservative SSU sequences (Fig. 1). Sequences from S. macrophylla (KY996315) appeared in a clade including S. caprifici and S. superbus. ...
... However, the sequences of F. richardi sp. n. were identical with those of a nematode identified as F. altermacrophylla (Lloyd & Davies, 1997) Davies & Bartholomaeus, 2015. This nematode was collected from Ficus rubiginosa growing near the Opera House in Sydney, NSW, Australia, in April 1999 (WNC 2290). ...
Ficophagus from collecting trips in eastern Australia, made over 15 years, are summarised and show that species of the genus occurred widely in sycones of Ficus , subgenus Urostigma , section Malvanthera . Two new species (based on morphological differences and molecular sequencing) are described: Ficophagus elizabethae sp. n. from Ficus macrophylla , F. rubiginosa and F. obliqua , and Ficophagus richardi sp. n. from Ficus obliqua ; and a morphospecies, Ficophagus Morphospecies malandicus from Ficus obliqua. Ficophagus elizabethae sp. n. is characterised by having the excretory pore (EP) opening from the level of the junction of the conus and shaft of the stylet to that of the knobs, a relatively long procorpus (1.0-2.5 times length of stylet), female tail with an obliquely truncate tail with a hyaline area and a finely to broadly rounded tip which may be mucronate; post-vulval uterine sac (PUS) ca one vulval body diam. (VBD) in length; rose-thorn-shaped spicule with distinct rostrum and prominent condylus; and genital papillae arranged as largest pair adcloacal, second pair posterior to mid-tail length, and third small pair near tail tip; and was collected from Sydney in New South Wales, to Bundaberg in Queensland (QLD). Ficophagus richardi sp. n. is characterised by having the EP opening at the level of the junction of the stylet shaft and conus, a labial cap which is raised around the opening for the stylet; procorpus 0.8-1.7 times length of the stylet, PUS <1 VBD in length, long uterus, and female tail with a V-shaped hyaline area at the bluntly rounded tip; rose-thorn-shaped spicule with a small rostrum and prominent condylus, three pairs genital papillae, first and largest on anterior cloacal lip, second at 70% of tail length measured from cloacal aperture, and third near tip, and was collected from Ban Ban Springs in the south to the Bundaberg region in the mid-north of QLD. In addition, in the absence of pertinent molecular sequences, a morphospecies is described. Ficophagus Morphospecies malandicus is characterised by having the EP opening anterior to the junction of the stylet conus and shaft, procorpus 0.9-2 times length of stylet, a short PUS usually <1 VBD long, short uterus, rose-thorn-shaped spicule with a raised condylus and prominent rostrum, and three pairs of subventral papillae on the tail (one adcloacal, one posterior to mid-tail and one near tail tip); and was collected from the Atherton Tableland, QLD. A table comparing morphological characteristics is provided to help with identification of Ficophagus nematodes from figs of the section Malvanthera in eastern Australia.
... However, in Schistonchus the character does not seem clearly correlated with the phylogenetic status. For example, the PUS is relatively short in S. caprifici and S. superbus but long in S. macrophylla, forming a neighboring clade with these two species (Lloyd & Davies, 1997;Zeng et al., 2013;Davies et al., 2015Davies et al., , 2020a, and short in S. pumilae n. sp. and S. hirtus, but rather long in S. cassowaryi, which is typologically similar to these two (Zeng et al., 2010;Davies et al., 2013b) (Fig. 1). Thus, the character could be closely tied to its function and evolving rapidly in each species. ...
... In the other nominal Schistonchus s.s. species, S. caprifici, S. m. macrophylla, S. m. lightfooti and S. flagellobenghalensis, and in the species inquirendae, S. osmani and S. racemosa, sperm has been described as 'flagellate' in the male testis/vas deferens and/or the spermatheca of females (Gasparini, 1864;Reddy & Rao, 1986, Lloyd & Davies, 1997Anand, 2002;Bajaj & Tomar, 2014;Davies et al., 2020a). By contrast, S. hirtus, S. superbus, S. athertonensis, S. molochi, S. cassowaryi and S. mucroracemosus have been described with amoeboid sperm (Zeng et al., 2010(Zeng et al., , 2013Davies et al., 2013aDavies et al., , 2013bBajaj & Tomar, 2014). ...
A Schistonchus species was isolated from the syconia of the creeping fig, Ficus pumila , collected from Miyazaki, Japan. The nematode was considered an undescribed species based on its typological characters, molecular profiles (near full length of small subunit and D3 expansion segment of large subunit of RNA) and phylogenetic status inferred from these two loci. The new species is characterised by the presence of flagellate sperm, spicule possessing small condylus, triangular rostrum with bluntly pointed tip and clear dorsal and ventral limbs, structure and arrangement of male caudal papillae, i.e. , presence of papilliform P3 and P3a and small and gland opening-like P4 (glandpapillae), relatively short post-uterine sac and female tail forming elongate conoid. It is phylogenetically close to S. hirtus . However, the new species is distinguished from other nominal Schistonchus species by its female tail shape forming elongate conoid, the structure and arrangement of male caudal papillae and several other typological characters. In addition, the host fig of the nematode, F. pumila , is a creeper fig species belonging to subgenus Synoecia (section Rhizocladus ; subsection Plagiostigma ), from which no fig-associated nematode has been reported so far, i.e. , the present study is the first report of Schistonchus (and other syconia-inhabiting nematodes) from the Ficus subgenus Synoecia .
... Following taxonomic revision made by Davies ((Davies et al., 2015)) North American species previously referred to as Schistonchus are now identified as Ficophagus. (Jauharlina et al., 2015;Davies et al., 2015;Gupta et al., 2021;Reddy and Rao, 1986;Krishnan et al., 2010;Kolaei et al., 2016;Kanzaki et al., 2023;Bajaj and Tomar, 2014;Zeng et al., 2007Zeng et al., , 2010Zeng et al., , 2011Zeng et al., , 2013aZeng et al., , 2013bZeng et al., , 2013cAnand, 2002;Fard and Zare, 2020;Rajeshwarianand, 2002) ( Bartholomaeus et al., 2009Bartholomaeus et al., , 2012Lloyd and Davies, 1997;Davies et al., 2010Davies et al., , 2013aDavies et al., , 2013bDavies et al., , 2020bZhao et al., 2015) ( Kruger et al., 2021;Martin et al., 1973;Vovlas et al., 1998) ...
... The fig nematode, Ficophagus religiosus (Bajaj & Tomar, 2014) Davies & Bartholomaeus, 2015 in Davies et al., 2015, was initially characterised from sycones of Ficus religiosa from Haryana, India. The Schistonchus sensu lato (s.l.) are classified into three genera: Ficophagus, which comprises 30 species, Schistonchus containing 14 species, and Martininema, four species (Gasparrini, 1864;Kumari & Reddy, 1984;Reddy & Rao, 1986;Lloyd & Davies, 1997;Vovlas et al., 1998;DeCrappeo and Giblin-Davis, 2001;Anand, 2002;Zeng et al., 2007Zeng et al., , 2010Zeng et al., , 2011Zeng et al., , 2013aZeng et al., ,b,c, 20192020;Bartholomaeus et al., 2009Bartholomaeus et al., , 2012Davies et al., 2010Davies et al., , 2013aDavies et al., ,b, 2015Davies et al., , 2017aBajaj & Tomar, 2014;Zhang et al., 2019;Zhao et al., 2015, 2020, Gupta et al., 2021Zhao et al., 2022) (Bajaj & Tomar, 2014;Davies et al., 2015Davies et al., , 2017aGupta et al., 2021;Waghmare et al., 2022) have been described, mostly based on morphological and morphometrical features. Ficophagus glomerata Gupta, Tahseen & Borges, 2021, a new species, has been described by providing morphological and molecular information from Ficus racemosa in Karnataka, India. ...
A population of Ficophagus was discovered in the syconia of sacred fig (Ficus religiosa) grown in New Delhi (India). Based on morphology, morphometric characters and molecular data, we redescribed the population as Ficophagus religiosus. Here, major morphological characteristics are described in depth, with measurements of the male-female specimens’ taxonomic characters and photomicrography. Ficophagus religiosus can be easily identified by the spiral to J-spiral-shaped body, labial disc not offset, SE pore close to the base of the lip region, stylet length 17.7 (16-19) μm in males, and 19 (15-22) μm in females, a short post-uterine sac
of length 8.3 μm (5-10 μm), a spicule with broad and high condylus, capitulum flat or depressed, a small bluntly rounded rostrum, the spicule tip (bluntly rounded or pointed), no cucullus, no gubernaculum, three pairs (P3, P3a, P4) of subventral caudal papillae, and
broadly or bluntly rounded tail tip, without mucron. Additionally, new sequence data for the D2-D3 region of 28S rDNA (LSU) and 18S rDNA (SSU) marker genes are provided. Maximum likelihood and Bayesian methods were used to infer phylogenetic relationships
of the Indian population of F. religiosus with other Ficophagus species. Phylogenetic analyses based on D2-D3 and SSU molecular markers resulted in a clear separation of F. religiosus from other Ficophagus species. The present study redescribes the species F.
religiosus and provides molecular data to identify and establish phylogenetic relationships with other species.
... Ficophagus, which comprises 28 species, Schistonchus 14 species, and Martininema four species (Gasparrini 1864;Lloyd & Davies, 1997;Vovlas et al., 1998;DeCrappeo & Giblin-Davis, 2001;Zeng et al., 2007Zeng et al., , 2010Zeng et al., , 2011Zeng et al., , 2013aZeng et al., , b, c, 2019Zeng et al., , 2020Bartholomaeus et al., 2009Bartholomaeus et al., , 2012Davies et al., 2010Davies et al., , 2013aDavies et al., , b, 2015Davies et al., , 2017aBajaj & Tomar, 2014;Zhang et al., 2019;Zhao et al., 2015Zhao et al., , 2020Gupta et al., 2021). In India, five species of Ficophagus (Ficophagus antherobenghalensis, F. cuculloracemosus, F. mucrobenghalensis, F. religiousus, and F. glomerata) have been described, mostly based on morphological and morphometrical characterisation (Bajaj & Tomar, 2014;Gupta et al., 2021). ...
A population of Ficophagus was recovered from white fig ( Ficus virens ) in New Delhi, India. We further described the population as Ficophagus virens based on morphological and morphometric characters, and molecular data. A detailed description of key morphological features, measurements of taxonomic characters, and photomicrography of the male and female specimens are given here. The study also included additional parameters such as lip diam. and height, conus stylet, shaft stylet, knobs diam. and height, vulva position from anterior end, length and width of spermatheca, anal/cloacal body diam., vulval width, and ovary/testis length for better characterisation of species. In addition to 28S rDNA (D2/D3), new sequence data from small subunit rDNA (18S) and mitochondrial cytochrome oxidase I COI marker gene were added. The D2/D3 sequence of F. virens was most similar to the sequence available for the Australian population of F. virens in GenBank. Maximum likelihood (ML) and Bayesian methods were used to analyse phylogenetic relationships of the Indian population of F. virens with those of the Australian populations and other Ficophagus species. This species is a new record from Delhi, India, and hence this report provides a new geographical location for the F. virens nematode after the first report from Australia.
A new species of the genus Ficophagus was recovered from the syconia of Ficus pisocarpa from Xishuangbanna, Yunnan province, China. It is described herein as Ficophagus pisocarpae n. sp. and is characterised by possessing the combined characters of a short PUS (6.4-9.3 μ m or 0.3-0.4 VBD long), an excretory pore near the head, presence of crustaformeria, amoeboid sperm, three pairs of subventral papillae on the male tail (one pair just adcloacal (P2), one pair halfway between cloacal aperture and tail terminus (P3a), and one pair near tail tip (P4)), rounded male tail tip without mucron, absence of gubernaculum and rose-thorn-shaped spicule with conical rostrum. Ficophagus pisocarpae n. sp. was differentiated from other sequenced species by the partial small subunit (SSU) rRNA gene and D2-D3 expansion segments of the large subunit (LSU) rRNA gene. Phylogenetic analysis with the LSU D2-D3 expansion segment sequence suggested that F. pisocarpae n. sp. is clustered in the same highly supported monophyletic clade with F. annulatae , F. benjamina , F. curtipes and F. microcarpus . It differs morphologically from these species in lateral incisure number, EP position, spicule and uterus morphology and some morphometric characters.
Nematodes, including plant-parasitic nematodes, are ubiquitous in all niches of forest ecosystems, but very rarely they have been implicated in forest mortality, forest decline, or losses of productivity. However, these problems are most likely underdiagnosed due to the concealed nature of these organisms, as well as the unspecific damage to the hosts, and complex species interaction. This task becomes even more complicated and economically unfeasible due to large spatial scales, remote locations and harsh terrain, and long times for economical return characteristic of forests. In the present review, existing information on abundance, composition, and dynamics of nematode communities, as well as pathogenic species, associated with the most representative groups of trees of the main forest types was compiled. The objective of this review was to find general patterns and key examples for each tree/forest type on the two major communities of nematodes associated with forest trees: soil-dwelling nematodes and stem-/canopy-dwelling nematodes, which are mostly transmitted to the aerial parts of the trees through insect vectors. This chapter offers relevant information on nematode infestation in forests and suggestions on their sustainable management.
A population of Schistonchus caprifici (Gasparrini, 1864) Cobb, 1927 was recovered from syconia of Ficus palmata grown in New Delhi (India). The population was identified based on morphological and molecular characteristics. The morphological and morphometric features of the Schistonchus population from Ficus palmata were found to be consistent with the original and subsequent descriptions of S. caprifici. This comprehensive morphological study of S. caprifici from India added additional information on morphological and morphometrical characters such as lip shape, stylet, vulva position from anterior end, spermatheca, anal/cloacal body diameter, vulval structure, and ovary/testis length. Alongside some morphological descriptions including spicule shape and gubernaculum in males, vulval lips and vaginal lines, post uterine sac, tale shape, and terminus are amended. The identification of this population was further confirmed by using sequences of the D2-D3 region of 28S rDNA (LSU), 18S rDNA (SSU), and COI of mt DNA, and its phylogenetic relationship with the S. caprifici from Turkey, Spain, Italy, and other species from around the world was established. A tabular key for the species under the genus Schistonchus is provided.
A new species of the genus Ficophagus was recovered from the syconia of Ficus variegata from Shenzhen, Guangdong province, China. It is described herein as Ficophagus giblindavisi n. sp. and is characterised by possessing the longest stylet in males (35.1-45.8 μ m) and most lateral incisures (5) of all currently described species in the genus, a short PUS (8.4-11.4 μ m or 0.3 VBD long), excretory pore situated at or posterior to the nerve ring, amoeboid sperm, three pairs of subventral papillae on the male tail, rounded male tail tip with a mucron, absence of gubernaculum and sickle-shaped spicules with a terminal cucullus. Ficophagus giblindavisi n. sp. was differentiated from other sequenced species by the partial small subunit (SSU) rRNA gene and D2-D3 expansion segments of the large subunit (LSU) rRNA gene. Phylogenetic analysis with the LSU D2-D3 expansion segment sequence suggested that F. giblindavisi n. sp. is clustered in the same highly supported monophyletic clade with F. auriculatae and F. fleckeri .
Studies were conducted in Italy on the biology of Schistonchus caprifici (Nemata: Aphelenchoididae). This nematode parasitizes and reproduces on caprifig (Ficus carica sylvestris) inflorescences and also in the haemocoel of the fig pollinator wasp Blastophaga psenes. Staminate and pistillate florets of caprifig infected by S. caprifici exhibited necrosis and cavities in the cortical parenchyma of floret peduncles and stamen filaments. Schistonchus caprifici attacked pistillate florets of commercial fig (F. carica); however, S. caprifici population densities were less (P= 0.05) in commercial fig than in caprifig (550 nematodes vs. 12,000 nematodes/g fresh floret tissues, respectively). Schistonchus caprifici was spread from infected to healthy caprifig inflorescences through oviposition behaviour of the wasp, Blastophaga psenes. Wasp development occurred in inflorescences concomitantly infected with nematodes, which colonized and started reproducing in the haemocoel of wasp larvae. Schistonchus caprifici persisted in the pupae and winged females that developed in the pistillate florets. Wingless male wasps were not infected by the nematode. Commercial fig inflorescences were not receptive to wasp development, although these inflorescences were visited by ovipositing wasps which vectored nematodes in their florets. Nematode infection may result in premature inflorescence decay. There was no evidence of any adverse effect on B. psenes due to S. caprifici infection. The association between S. caprifici and B. psenes on caprifig can be defined as a parasitic relationship because of nematode reproduction in the wasp's haemocoel. Nematode populations from wasps and caprifig florets did not differ in their morphology.
A neotype of Aphelenchus avenae is designated and the variation shown by the species described. Metaphelenchus rhopalocercus and M. micoletzkyi are synonyms of A. avenae, and M. sacchari is species inquirenda.