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Photographs and SEM micrographs of harvestmen from the families Kimulidae, Manaosbiidae, and Samoidae. A. Dorsal view of undescribed species 1 (Kimulidae). B. Lateral view of eye mound and dorsum of kimulid sp. 1. C. Dorsal view of undescribed species 2 (Kimulidae). D. Dorsal view of undescribed species 3 (Kimulidae). E. Lateral view of kimulid sp. 3. F. Dorsal view of female Rhopalocranaus albilineatus (Manaosbiidae) with typical dorsal pattern. G. Dorsal view of male R. albilineatus showing unusual pattern (white rings around several dorsal spines) exhibited by individuals captured from Salybia. H. Dorsal view of a male Cranellus montgomeryi (Manaosbiidae). I. Dorsal view of a female C. montgomeryi. J. Tarsus of leg I of a male C. montgomeryi showing spindled morphology. K. SEM micrograph of distitarsus of leg I of female C. montgomeryi, scale bar = 250 μm. L. Dorsal view of Maracaynatum trinidadense (Samoidae). M. Dorsal view of Pellobunus longipalpus (Samoidae). N. SEM micrograph of the dorsal view of unidentified species collected from bromeliads (Samoidae), scale bar = 0.5 mm. O. SEM micrograph of chelicerae and pedipalps of unidentified samoid, scale bar = 250 μm. P. SEM micrograph of the ventral surface of unidentified samoid, scale bar = 0.5 mm.
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From 2005-2007, we observed 20 species of harvestmen representing nine families, primarily in northern Trinidad. Of these taxa, we collected previously unreported or undescribed species for the families Kimulidae, Samoidae, Sclerosomatidae, and Stygnommatidae. In this paper, we provide a brief summary of the characters and natural history,
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Gamelia bennetti sp. nov. is described from Trinidad, Trinidad and Tobago, West Indies, and compared with members of the Gamelia abas species group: G. abas (Cramer, [1775]), G. berliozi Lemaire, 1967, G. lichyi Lemaire, 1973, G. rubriluna (Walker, 1862) and G. septentrionalis (Bouvier, 1936). A photographic record suggests G. bennetti sp. nov. may...
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... Harvestmen adalah haiwan omnivor yang makan haiwan invertebrata, tumbuhtumbuhan, serta kulat yang reput dan boleh dijumpai di semua ekosistem daratan seperti di bawah batu, atas pokok, pada kayu buruk dan atas tanah (Acosta & Machado 2007;Schaus et al. 2013;Townsend et al. 2008). Kelimpahan komuniti harvestmen di habitat berhutan lebih tinggi berbanding habitat terbuka kerana factor abiotik yang lebih lembab dan terlindung untuk mengelakkan kehilangan air (De Poel & De Smedt 2016). ...
... Kajian-kajian lepas ke atas harvestmen adalah tertumpu kepada kajian taksonomi (Townsend et al. 2008), filogeni melalui pendekatan molekul (Hedin et al. 2012;Oberski et al. 2018) dan morfometrik (De Bivort et al. 2010). Majoriti kajian lepas tentang harvestmen dijalankan luar dari Malaysia (Černecká et al. 2017;Cook et al. 2013;Garwood et al. 2011). ...
Taxonomy-based morphometric study is a technique often used to separate species. This study aimed to determine discrimination among species of harvestmen based on morphometric analysis. Harvestmen samples collected from Lata Jarum, Pahang deposited at the Centre of Insect Systematic, National University of Malaysia (CISUKM) were used. A total of 21 morphometric parameters was measured using the Dino-Lite Digital Microscope in millimeter (mm) unit. Result from univariate analysis were unable to distinguish between the species but analysis from Canonical Discriminant Analysis (CDA) using the Ratio of One could clearly distinguish the species into four clusters namely: Zaleptus quadrimaculata with Dentobunus luteus, Marthanella ferruginea, Marthana sp.1 and Marthana sp.2. Principal Component Analysis (PCA) using Ratio of Two could only distinguish species into two clusters: Cluster 1 consists of Zaleptus quadrimaculata, Dentobunus luteus, Marthana sp.1, Marthana sp.2 and Cluster 2 only Marthanella ferruginea. The important parameter detected by CDA were EyeW/MaxLDS, DBEye/MaxLDS, FemurL/MaxLDS and TibiaL/MaxLDS while important parameter detected by PCA were EyeW/MaxPro, DBEye/MaxWPro and MaxLOpi/MaxWOpi. In conclusion, morphometric studies can be used in identification of harvestmen species and can provide early information to avoid misidentification of species in future. ABSTRAK Kajian taksonomi berasaskan morfometrik merupakan satu teknik yang sering digunakan untuk pemisahan spesies. Kajian ini bertujuan untuk mengkaji perbezaan antara spesies harvestmen berasaskan analisis morfometrik. Sampel harvestmen yang telah dikumpul dari Lata Jarum,
... In addition, we compared the surface morphology of the chelicerae and pedipalps of C. marginalis with those of two additional cosmetid harvestmen, Paecilaema inglei Goodnight & Goodnight, 1947 and Erginulus clavotibialis (Pickard-Cambridge, 1905). In the past, we have handled over 100 live individuals of each species (Townsend et al. 2008;Schaus et al. 2013). While we have never observed stridulation by E. clavotibialis, we have infrequently detected vibrations emanating from adult P. inglei. ...
Stridulatory organs have not been previously investigated for harvestmen in the family Cosmetidae. During a field study, we observed the infrequent production of vibrations by adult Cynorta marginalis Banks, 1909. Using SEM, we examined the surfaces of several appendages for potential stridulatory organs. Our observations indicate that C. marginalis has denticles on the mesal surfaces of the basichelicerites that when rubbed together may function as an isomorphous stridulatory organ. In addition, there are denticles on the ectal surfaces of the basichelicerites and furrowed ridges on the femora of the pedipalps that may represent heteromorphous stridulatory organs. We did not observe any sexual dimorphism in morphology at either anatomical location. We also examined the appendages of two additional cosmetid harvestmen: Paecilaema inglei Goodnight & Goodnight, 1947, a species that also stridulates when held and Erginulus clavotibialis (Pickard-Cambridge, 1905), a species that has not been observed to produce vibrations. As in C. marginalis, we observed denticles on mesal and ectal surfaces of the basichelicerites and a furrowed ridge on the mesal surfaces of the femora of the pedipalps of adult P. inglei. In contrast, the basichelicerites of E. clavotibialis had relatively fewer and smaller denticles on the external surfaces of the chelicerae and the mesal surface of the femora of the pedipalps lacked ridges and were relatively smooth. Our comparative morphological data supports the hypothesis that there are cosmetid harvestmen that may use surface features on the chelicerae and pedipalps to produce vibrations which may function as a secondary defense mechanism.
... Voucher specimens of these specimens will be deposited in the collections of the American Museum of Natural History (AMNH) and the Universidad de Costa Rica. Adult females of R. albilineatus and P. calcariferus were collected as part of a series of field surveys and natural history studies from 2005 to 2008 in Trinidad, W.I. (locality information is detailed in Townsend et al. 2008). These specimens were used in the study by Bennett & Townsend (2013) to describe interspecific variation in the microanatomy of the ovipositor. ...
In many species of scorpions (Order Scorpiones) and spiders (Order Araneae), mating plugs (gel-like or sclerotized) are deposited in the female reproductive tract, minimizing the possibilities of further matings. Although harvestmen (Order Opiliones) have direct sperm transfer via copulatory organs (penis or spermatopositor in the male and ovipositor in the female), prior studies of the reproductive anatomy of these arachnids have not reported the occurrence of these structures. With the aid of scanning electron microscopy, we observed distinct masses of amorphous, gel-like material in the distal openings of the ovipositors of six different specimens including representatives of the families Cosmetidae (Cynorta marginalis and Flirtea picta), Gonyleptidae (Discocyrtulus bresslaui and Geraeocormobius sylvarum), Kimulidae (Kimula sp.) and Metasarcidae (Metasarcus clavifemur). Dissection of one of these structures from a specimen of the cosmetid harvestman C. marginalis revealed a detailed endocast of the internal surfaces of the female reproductive tract. Considerable interspecific variation in the relative sizes and shapes (smooth and rough) of the amorphous gel-like structures were observed, but evidence of sclerotized components (i.e., parts of the male genitalia) was not found. The location and morphology of the gel-like structures are consistent with observations of mating plugs in other arachnids. The composition, source (male, female or both) and function of these amorphous gel-like structures (i.e. effective matting plug) requires additional study.
... We compared the morphology of different podomores (trochanter, femur, patella, tibia, and tarsus) from the pedipalps of nymphs and adults of three relatively common species of Neotropical cosmetid harvestmen using specimens that had been collected in previous studies (Townsend, Proud, & Moore, 2008;Townsend, Schaus, Zvonareva, Illinik, & Evans, 2017;Townsend et al., 2009;Walker & Townsend, 2014). Individuals of Cynortula granulata Roewer 1912 ( Figure 1 to 24 August 2015. ...
... These species include Cranellus montgomeryi Goodnight and Goodnight, 1947 1947) (Stygnidae). Details for collections sites and dates for these specimens are described in Townsend et al. (2008) and Rodriguez, Townsend, Johnson, et al. (2014). ...
In arachnids, pedipalps are highly variable appendages that may be used in feeding, courtship, defense, and agonistic encounters. In cosmetid harvestmen, adults have pedipalps that feature flattened femora, spoon-shaped tibiae, and robust tarsal claws. In contrast, the pedipalps of nymphs are elongate with cylindrical podomeres and are adorned with delicate pretarsi. In this study, we used scanning electron microscopy to examine the distribution of cuticular structures (e.g., sensilla chaetica, pores) on the elements of the pedipalps of adults and nymphs of three species of cosmetid harvestmen. Our results indicate that there is considerable ontogenetic variation in the morphology of the trochanter, femur, patella, tibia, and tarsus. The pretarsus of the nymph has a ventral patch of setae that is absent from the adult tarsal claw. We observed this structure on all three cosmetid species as well as on the pedipalps of an additional seven morphospecies of nymphs collected in Belize and Costa Rica. This structure may represent a previously unrecognized autapomorphy for Cosmetidae. Examinations of the pedipalps of antepenultimate nymphs of additional gonyleptoidean harvestmen representing the families Ampycidae, Cranaidae, Manaosbiidae, and Stygnidae revealed the occurrence of unusual, plumose tarsal setae, but no setal patches on the tarsal claw.
... Adult Cynortula granulata Roewer, 1912 were collected in July and August 2005-2006 from forested habitats in Trinidad, W.I. We captured harvestmen (n 5 158) by hand from multiple locations throughout the Northern Range and central Trinidad (descriptions of habitats and specific locations are provided in Townsend et al., 2008). All harvestmen were placed in 70% ethanol or 10% buffered formalin. ...
... Additional behavioral studies are needed to assess whether males employ secondary defenses differently than females or whether potential predators specifically target the legs of adult males. In the forests of Trinidad, C. granulata is a habitat generalist and is most commonly found in the leaf litter and beneath logs (Townsend et al., 2008;Proud et al., 2011). However, C. granulata also infrequently occupies perches on tree buttresses that are 1-2 m above the ground (Burns et al., 2007). ...
Previous studies of leg injuries in har-vestmen have focused on the fitness consequences for individuals that use autospasy (voluntary detachment of the leg) as a secondary defense mechanism. Leg damage among non-autotomizing species of laniatorean harvestmen has not been investigated. Under laboratory conditions, we damaged femur IV of Cynorta mar-ginalis and observed with scanning electron microscopy (SEM) the changes in these wounds over ten days. We also used SEM to examine leg damage from individuals of three species of cosmetid harvestmen that were collected in the field. On the basis of changes in the external surface of the hemolymph coagulum, we classified these wounds as fresh (coagulum forming), recent (coagulum with smooth surface), older (coagulum is scale-like with visible cell fragments), and fully healed (scale replaced by new cuticle growth on the terminal stump). Our observations indicate that wound healing in harvestmen occurs in a manner comparable to that of other chelicerates. Leg injuries exhibited interspecif-ic variation with respect to the overall frequency of leg wounds and the specific legs that were most commonly damaged. In addition, we measured walking and climbing speeds of adult C. marginalis and found that individuals with fresh injuries (lab-induced) to femur IV walked at speeds significantly slower than uninjured adults or individuals collected from the field that had fully healed wounds to a single leg. J. Morphol. 000:000–000, 2016. V C 2016 Wiley Periodicals, Inc.
... Based upon the detailed descriptions of Mora (1990), it does not appear that the female uses the ovipositor to probe the ground prior to egg deposition or to select specific sites within the mud nest for egg laying. With respect to reproductive morphology, Bennett and Townsend (2013) examined ovipositors of several individuals of the manaosbiid harvestman Rhopalocranaus albilineatus, a species that can be fairly abundant in the leaf litter and under logs ), but does not construct mud nests (Townsend et al. 2008). In this species, the ovipositor has four external lobes that are adorned with 2-3 peripheral setae that insert into undivided sockets. ...
The external anatomy of the ovipositor has generally been overlooked as a source of informative characters in systematic studies of laniatorean harvestmen. In this study, we used scanning electron microscopy to examine the ovipositors of nine species representing the families Manaosbiidae (five species) and Nomoclastidae (four species). Similar to the ovipositor morphology of many gonyleptoidean families, the distal tips of the ovipositors of these harvestmen have four external lobes, with the margins most commonly adorned with 10 large peripheral setae. In manaosbiid and nomoclastid species, these peripheral setae have undivided bases, striated shafts and undivided distal tips. There are typically three setae on each anterior lobe and two setae on each posterior lobe. The medial setae on both anterior and posterior lobes insert into sockets that are slightly more dorsal. We observed small, surface denticles, usually associated with a pore, on the external surface of the lobes. There was interspecific and intraspecific variation in the number and shape of these surface denticles. The association of pores with denticles on the ovipositor appears to be a feature common to not only both families but is also a trait that has not been observed on ovipositors in other families of laniatorean harvestmen.
... Adult C. anulata, P. calcariferus, and R. albilineatus were collected from forested habitats in northern and central Trinidad, West Indies from 2003-2008(Townsend et al., 2008). Individuals were captured by hand and preserved in 70% ethanol. ...
... Adult C. anulata, P. calcariferus, and R. albilineatus were collected from forested habitats in northern and central Trinidad, West Indies from 2003-2008(Townsend et al., 2008). Individuals were captured by hand and preserved in 70% ethanol. ...
... With respect to penis morphology, our descriptions of male genital morphology are similar to those previously published for these species Kury, 2003, 2011;Townsend et al., 2008Townsend et al., , 2011. In general, we found relatively little intraspecific variation in structures associated with the ventral plate or the glans penis. ...
Illustrations of penis morphology are essential components of species descriptions for harvestmen belonging to the suborder Laniatores. Male genitalia are important sources of taxonomic characters and are generally assumed to exhibit relatively little intraspecific variation. In contrast, descriptions of female reproductive morphology are rarely included in taxonomic descriptions of laniatorean harvestmen. As a result, relatively little is known about variation in the external features of the ovipositor. In this study, we used scanning electron microscopy to examine variation in male and female reproductive morphology among three species of harvestmen that are members of the superfamily Gonyleptoidea. Specifically, we examined the microanatomy of penises and ovipositors of Cynortula granulata (Cosmetidae), Phareicranaus calcariferus (Cranaidae), and Rhopalocranaus albilineatus (Manaosbiidae). Our results support the general observation that male reproductive morphology is conservative and displays little intraspecific variation. We observed considerable intraspecific variation in the number and shape of marginal setae on the ventral plate of the penis, but little or no variation in the morphology of the distal border of the ventral plate or the shape of the glans penis or stylus. With respect to female genitalia, we observed considerable intraspecific variation in the number of peripheral setae on the distal tip of the ovipositor. We also observed interspecific variation in the morphology of the peripheral setae (surface of the shaft and at shape of the distal tip), the distribution and morphology of smaller setae on the external surface of the ovipositor, and the surface texture of the external lobes. Our results indicate that there are several features associated with ovipositor morphology among laniatorean harvestmen that may represent potentially informative taxonomic characters. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.
... Individuals of some temperate sclerosomatid species are known to make vertical migrations-climbing on trees during the day and descending at dusk to forage on or near the ground [6,17,18]. In montane rainforests of Trinidad, two syntopic species belonging to the family Cranaidae often use fallen palm frond sheaths as diurnal shelter [22] or oviposition sites [45,54,55]. The use of the endocarp of a coconut as an oviposition site was also observed for a cranaid harvestman [56]. ...
In tropical rain forests, harvestmen assemblages are extremely diverse, with richness often exceeding 25 species. In the neotropics, there are published accounts of harvestmen faunas in South America rainforests (especially Amazonia), but relatively little is known about the community ecology of harvestmen in tropical forests of Central America. In this paper, we provide the first insights into the diverse assemblage of harvestmen inhabiting a wet forest at La Selva Biological Station, Costa Rica. Over five field seasons, we recorded 38 species. During our 2009 field season, we examined variation in species abundance, richness, and composition between adjacent successional forests (young secondary, mature secondary, and primary forests) as well as between distinct habitats (ground/litter layer and shrub/tree layer). Based on night samples (but not day), our results indicate that there are only minor differences in species composition and relative abundance between the forest ages, but no differences in richness. The ground/litter layer and shrub/tree layer habitats differed markedly in species composition, species richness, and relative abundance of several species. Our analysis of covariance supports the hypothesis that leg length is related to climbing behavior for several species belonging to Eupnoi and Laniatores.
... m Trinidad, and also attributed erroneously to a Bolivian city of same name. Roewer wrongly sexed P. cingulatus as male, which led him to pro-pose it in a different genus, even though he described it with a similar color pattern as P. calcariferus. Later, Goodnight and Goodnight (1947) recognized P. calcariferus from material collected in Trinidad. Townsend et al. (2008) stated that " male Phareicranaus calcariferus are smaller in total body size (7.4–9.6 mm) than Santinezia serratotibialis (9.6–11.9 mm) and also lack spines on ventral surface of coxae IV. Males S. serratotibialis have more prominent spines in femur IV " . Townsend et al. (2009) studied the postembryonic development of S. serratotibiali ...
A reanalysis of the species relationships in the genus Phareicranaus Roewer, 1913, here considered the senior synonym of Acanthocranaus Roewer, 1913, Santinezia Roewer, 1923 and Comboyus Roewer, 1943, is carried out, using information unavailable in a previous analysis (of Santinezia). Subfamilies Cranainae and Prostygninae may be not monophyletic, since the generic relationships among cranaines are still obscure; it seems that Phareicranaus is closely related to Ventrivomer Roewer, 1913 and Ventrisudis Roewer, 1963, based on presence of a ventral apophysis on male coxa IV and cylindrical pedipalpal femur. The analysis performed with equal weighting generated 64 equally parsimonius trees (275 steps, ci=.24; ri=.60), and the strict consensus tree differs substantially from previous analyses. Based on results of the equally weigthed cladistic analysis, Phareicranaus is supported by the following unambiguous synapomorphies: penis base of the glans ringed; pedipalpal femur with a stout dorsoapical spine; strong ventral tubercles on pedipalpal femur and; area II invading area I (the two halves do not touch each other). Two new species are described, Phareicranaus divisor (type-locality: Brazil, Acre, Parque Nacional da Serra do Divisor) and P. patauateua (type-locality: Brazil, Para, Ourem, Patauateua, fazenda Gaviao). Male genitalia are described for the following cranaids: Chiriboga albituber Roewer, 1959; Metacranaus tricalcaris Roewer, 1913; Phareicranaus calcariger (Roewer, 1913), comb. n.; P. albimedialis (Goodnight & Goodnight, 1943); P. festae Roewer, 1925; P. parallelus Roewer, 1925; P. angelicus (Roewer, 1963) comb. n.; and P. ortizi (Roewer, 1952) and Ventrisudis mira Roewer, 1963. The following new combinations are proposed: from Acanthocranaus, Phareicranaus calcariger (Roewer, 1913); from Santinezia, Phareicranaus albilineatus (Roewer, 1932); P. angelicus (Roewer, 1963); P. arthrocentricus (Mello-Leitao, 1943), P. biordi (Gonzalez-Sponga, 1991); P. calcarfemoralis (Roewer, 1917); P. calcaritibialis (Roewer, 1915); P. capayitaensis (Gonzalez-Sponga, 2003); P. contrerasi (Gonzalez-Sponga, 2003); P. circumlineatus (Gonzalez-Sponga, 1989); P. curvipes (Roewer, 1916); P. duranti (Gonzalez-Sponga, 1989); P. fronteriza (Gonzalez-Sponga, 2003); P. furva (Pinto-da-Rocha & Kury, 2003); P. giganteus (Roewer, 1913); P. glaber (Gonzalez-Sponga, 2003); P. gracilis (Pinto-da-Rocha & Kury, 2003); P. guaricoensis (Gonzalez-Sponga, 2003); P. heliae (Avram, 1983); P. hermosa (Pinto-da-Rocha & Kury, 2003); P. lamitisus (Gonzalez-Sponga, 2003); P. leonensis (Gonzalez-Sponga, 2003); P. lucifer (Pinto-da-Rocha & Kury, 2003); P. manauara (Pinto-da-Rocha, 1994); P. noctiscansor (Townsend & Milne 2010); P. onorei (Pinto-da-Rocha & Kury, 2003); P. ortizi (Roewer, 1952); P. parvus (Gonzalez-Sponga, 2003); P. sanarensis (Gonzalez-Sponga, 2003); P. simonbolivari (Avram, 1987); P. singularis (H. Soares, 1970); P. soledadensis (Gonzalez-Sponga, 2003); P. spinulatus (Goodnight & Goodnight, 1943); and P. subparamera (Gonzalez-Sponga, 2003). Santinezia sanarensis humocaroensis Gonzalez-Sponga, 2003, transferred to Phareicranaus, is elevated to species level, proposing the new combination and new status: Phareicranaus humocaroensis (Gonzalez-Sponga, 2003). Three new names are proposed: Phareicranaus roeweri to replace Nieblia festae Roewer, 1925, preoccupied by Phareicranaus festae Roewer, 1925; Phareicranaus goodnightorum to replace Santinezia magna Goodnight & Goodnight 1942, preoccupied by Phareicranaus magnus (Roewer, 1932); and Phareicranaus kuryi to replae Comboyus albilineatus Roewer, 1943, preoccupied by Santinezia albilineata Roewer, 1932, which is presently a junior synonym of Inezia curvipes Roewer, 1916 (now Phareicranaus curvipes). Phareicranaus cingulatus Roewer, 1932 and Santinezia serratotibialis Roewer, 1932 are newly synonymized with Goniosoma calcariferum Simon, 1879 (the type species of Phareicranaus). Additionally, Phareicranaus giganteus Roewer, 1932 is newly synonymized with Holocranaus longipes Roewer 1913, and the monotypic genus Meridia Roewer 1913 (type species M. palpalis Roewer 1913) is transferred from Manaosbiidae to Cranaidae, Cranainae.
... This is due, at least in part, to undersampling or lack of sampling the leaf litter, a microhabitat in which they can be relatively abundant (Kury 2007). Little is known about the natural history of these harvestmen, although Townsend et al. (2008a) provided observations concerning activity, habitat use and geographic distribution for Cranellus montgomeryi Goodnight & Goodnight 1947a and Rhopalocranaus albilineatus Roewer 1932 on the Caribbean island of Trinidad. The two basal segments on tarsus I of males in most species are generally enlarged and frequently fused (Kury 2007). ...
... Natural history.-Little is known about the natural history of harvestmen from the family Manaosbiidae. In Trinidad, Townsend et al. (2008a) reported that Rhopalocranaus albilineatus is a habitat generalist and exhibits an island-wide distribution. In contrast, Cranellus montgomeryi is a habitat specialist, with a distribution limited to montane rainforest and elfin woodland in the Northern Range. ...
In Central America, the family Manaosbiidae is recorded only from Panama and Costa Rica. Four species
occur in this region: Barrona williamsi Goodnight & Goodnight 1942, Bugabitia triacantha Roewer 1915, Poassa limbata Roewer 1943, and Zygopachylus albomarginis Chamberlin 1925. In this paper, we describe Barrona felgenhaueri new species (Cocle´ Province, Panama) and Bugabitia akini new species (Cocle´ Province, Panama) and report a new record for B. williamsi (Cocle´ Province, Panama). We used SEM to examine the penis morphology of Barrona Goodnight & Goodnight 1942 and the Caribbean species Cranellus montgomeryi Goodnight & Goodnight 1947 and Rhopalocranaus albilineatus Roewer 1932. We compared genital morphology of these species with published descriptions for Manaosbiidae from South America. With respect to genital morphology, we found that the most variable characters were the number and relative sizes of the setae that occur on the lateral margins of the ventral plate. Other features that exhibited interspecific variation included the shape of the ventral plate, the shape of the distal border of the ventral plate, and the shape and armature of the apex of the stylus.
