A Redescription and Family Placement of Buemarinoa Patrizii Roewer, 1956 (Opiliones, Laniatores, Triaenonychidae)
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Biologia Serbica, 2019, 41(1): 67-77 DOI 10.5281 /zenodo.3373487 Original paper A redescription and family placement of Buemarinoa patrizii Roewer, 1956 (Opiliones, Laniatores, Triaenonychidae) Ivo KARAMAN University of Novi Sad, Faculty of Sciences, Department of Biology and Ecology, Trg Dositeja Obradovića 2, 21000 Novi Sad, Serbia Received: 2 May 2019 / Accepted: 26 July 2019 / Published online: 21 August 2019 Summary. Buemarinoa patrizii Roewer, 1956, from a Sardinian Cave and Fumontana deprehendor Shear, 1977, from the Appalachian Region of eastern North America, are closely related and are the only representatives of the temperate Gondwanan family Triaenonychidae in the Northern Hemisphere. Based on their relevant characteristics, these two species, together with Flavonuncia pupila Lawrence, 1959, from Madagascar, are placed into a new tribus Buemarinoini tribus n., which represents an old phyletic lineage. It is a unique case of a phyletic lineage (tribus in this case) with three representatives present on three different continents. The elongated appendages in troglobite Buemarinoa patrizii are not treated as a troglomorphic trait. Elongation of appendages in troglobionts is not an adaptation, but a precondi- tion for settling in a subterranean environment. The main morphological differences of Fumontana deprehendor in comparision with Buemarinoa patrizii are treated as apomorhic. The unique terminal structure of the male copulatory apparatus, which includes a pair of flanking setose arms, a pair of glans glands and bifurcated sperm duct, define the Buemarinoini tribus n. Based on the male penis structure, Buemarinoini tribus n., shows closeness to a complex of spe- cies gathered around the genera Ceratomontia, Austromontia and Monomontia from South Africa and South America. The origin of Buemarinoini must be related to the areas of Northern Gondwana, from where Fumontana penetrated deeper into Laurasia. Lectotypes are designated for Buemarinoa patrizii type material. Key words: amphi-Atlantic, Laurasia, lectotype, new tribus, Northern Gondwana, troglomorphic. INTRODUCTION habitats of the Temperate Zone. They have survived in these areas mainly as relict cryptobionts. Subordo Laniatores is a highly species-rich A disproportional presence of basal groups of Laniato- group of harvestmen with nearly 4,200 described res between Temperate Zones of the Southern and North- species (Kury 2013). Its representatives are most ern Hemisphere is evident. Basal groups of Laniatores in common in the tropics, where they display the the Temperate Zones of Northern Hemisphere include rep- highest level of species diversity. Cyphophthalmi resentatives of the superfamily Travunioidea (Travuniidae, and Palpatores, the remaining two suborders of Cladonychiidae, Paranonychidae and Cryptomastridae), harvestmen, show a somewhat greater diversity the family Phalangodidae and, symbolically, the family Tri- outside the tropics. As for Palpatores, with the ex- aenonychidae; while in the Southern Hemisphere they are ception of the family Sclerosomatidae, its overall represented only by the family Triaenonychidae. Such a pic- diversity is associated with Temperate Zone areas. ture is suggestive, and (among other factors) illustrates the The three suborders share the fact that their basal great age of the family Triaenonychidae. The presence of the groups (early-diverging lineages) are present almost basal groups of subordo Palpatores shows a similar pattern exclusively outside of the tropical belt, in cryophilic of presence, exclusively in habitats of the Temperate Zone. E-mail: [email protected] I. Karaman Fig. 1. Buemarinoa patrizii, paralectotype female 1.4 mm, side view. (Scale bar = 0.5 mm.) Buemarinoa patrizii Roewer, 1956, a little-known cave 1), I observed that it is actually a member of the family Tri- species from Sardinia after its description was mentioned in aenonychidae, close to the species Fumontana deprehendor a few publications, as one of the European representatives Shear, 1977 (Fig. 2), from the Appalachian Region in North of the family Travuniidae (Martens 1978; Kury and Mendes America, which is the only representative of this Gondwa- 2007; Kury et al. 2014), in which Roewer originally placed it. nan family in the Northern Hemisphere. The unique char- Recently, Derkarabetian et al. (2018) classified this species acteristics of these two species are reflected in the specific in the Laurasian family Cladonychiidae. Kury and Mendes and complex structure of the male copulatory apparatus. A (2007) expressed doubts about the status of the genus Bue- similar penis structure is present in the species Flavonuncia marinoa, but not about its family affiliation. In all of these pupila Lawrence, 1959, from Madagascar. All three species important monographs and studies, which mentioned or based on this important and most indicative character stand briefly referred to B. patrizii, a mistake has clearly been in contrast to all other species in the family Triaenonychidae. made, because it was not possible to recognize the family The family Triaenonychidae is not clearly defined. The affiliation of the species without insight into the actual speci- reasons for this have been elaborated in detail by Hunt and mens. Indeed, the original description of this species did not Hickman (1993), and have been attributed to the fact that significantly differentiate it from other travuniids, as explic- it is an extremely old group and an old diversification. Al- itly stated in the diagnosis of the genus. Also, a significant most all species of Triaenonychidae have limited distribution, mistake in the description, where the typical triaenonychid which best illustrates their relic character. type of the posterior leg claws in this species was listed as Actual division of the family is based on the structure peltonychium type, appears to have misled the above listed of the sterna and tarsal claws of the hind legs, homoplastic authors. The great taxonomic significance that was previously characters which are variable at all taxonomic levels. Because attributed to this character could have been earlier identified, of this, many authors expressed doubts about the validity of if it had been correctly interpreted, enabling experts to study the division (Forster 1954; Maury 1988; Hunt and Hickman this species in more details. In addition to these missteps, 1993). However, division of the family into four subfamilies placement with the above mentioned families was considered is still in use (Kury et al. 2014; Kury 2018): Triaenonychi- to be “logical”, given the geographical position of Sardinia. nae Sørensen, 1886 (South Africa, Madagascar, New Zea- This must have had a significant effect on speculations con- land, Tasmania, Australia, South America, New Caledonia, cerning species placement, which failed to take into account Eastern North America); Adaeinae Pocock, 1902 (South the highly complex biogeography of the Mediterranean and Africa and Western Australia); Triaenobuninae Pocock, its limitrophic regions. 1902 (Tasmania, Southeast Australia, New Zealand, South Examining type specimens of Buemarinoa patrizii (Fig. America, Madagascar) and Soerensenellinae Forster, 1954 68 Biologia Serbica 41 A redescription and family placement of Buemarinoa patrizii Roewer, 1956 (Opiliones, Laniatores, Triaenonychidae) Fig. 2. Fumontana deprehendor, male 1.65 mm, side view. (Scale bar = 0.5 mm.) (New Zealand and South Africa). Species of the New Zealand MATERIALS AND METHODS monotypic family Synthetonychiidae differ from Triaenony- chidae by their unusual appearance, but not in other relevant Photographs of habitus morphology as well some de- characters. Therefore, some authors have expressed doubts tails (pedipalps, ocular tubercle) were taken using a Leica about the validity of its existence (Martens 1986; Hunt and M205C stereo microscope equipped with a Leica DFC290 Hickman 1993). HD digital camera. Final images were taken at different focal A comprehensive revision of Triaenonychidae is clearly planes and combined using Leica Application Suite V4.11 necessary, in order for species of this family to be grouped software. Photographs from slides were taken using a Zeiss into subfamilies that are based on real phyletic relation- Axio Imager A1 compound microscope equipped with an ships and valid characters. Unfortunately, this is currently AxioCam MRc5 digital camera. Final images were taken not the case. Therefore, I am placing genera Buemarinoa, at different focal planes and combined using Helicon focus Fumontana and Flavonuncia provisionally into the sub- stacking software. Drawings were made in Adobe Illustrator family Triaenonychinae, as nominal one of the established CS2 on a Genius EasyPen M610X graphics tablet. Dissected subfamilies, and because Fumontana and Flavonuncia had parts were mounted on slides in glycerol. After glycerol the been already placed in it by their authors. To emphasize the ovipositor and penises were temporary mounted on slides in phyletic proximity of these three geographically very distant Faure’s mounting medium (40 g chloral hydrate, 10 g gum genera, I decided, although it is not common practice in the Arabic; 5 g glycerin, 5 g glacial acetic acid, 5 g glucose, and classification of Opiliones, to establish for them a new tribe distilled water). - the Buemarinonini tribus n. For comparison a male specimen of Fumontana dep- rehendor from USA (Tennessee, Cocke Co, road to Cosby Campground, 27. 08. 2005, leg. M. Hedin et al.) has been