Belongs within: Neopilionidae.
The Ballarrinae is a group of very small long-legged harvestmen found in Australia, southern Africa and South America. Ballarrines are characterised by the possession of elongate pedipalps with an extremely long patella, retroflexed tibia and reduced terminal claw on the tarsus. In the Australian genera Ballarra, Plesioballarra and Arrallaba, a barbed process is present on the left side of the penis at the junction between shaft and glans; such a process is absent in the South American Americovibone lanfrancoae (the male of the South African Vibone vetusta has not been described).
New Zealand fills a biogeographical gap
Published 28 March 2012
New paper, and new species of phalangioid harvestman, out (Taylor 2016)! And one that I’m pretty excited by, even if the vagaries of time allocation mean that I haven’t been able to get the post out until a couple of weeks after it happened. After several years of studying New Zealand’s long-legged harvestman fauna, I have to confess I was getting a bit complacent about. I certainly knew that I had not seen every species that the country had to offer, but I still thought that there were no real surprises remaining. The overall outline had become clear; any species of long-legged harvestman remaining to be described from New Zealand would be fairly closely akin to those already known.
Oh boy, was I wrong.
At some point last year (or maybe the year before), I was sorting through a jarful of specimens that were still waiting on my attention. In one of the vials, its contents collected in a remote part of the south-west South Island, was a tiny, wispy specimen that I at first glance paid little mind to. Newly-hatched juveniles are not uncommonly collected; they are almost always unidentifiable and end up being just chucked back into the jar never to be looked at again. Nevertheless, I pulled the specimen out to confirm that my first impression was correct. I placed the specimen in a dish under the microscope and glanced through the eyepiece. Then looked again, my eyes doubtless boggling. I may have even sworn a little. Not only was the specimen not a juvenile but fully adult, it was something I had long given up on seeing from New Zealand: a ballarrine.
The Ballarrinae are an unusual group of harvestmen that were not recognised until fairly recently. The group was named by Hunt & Cokendolpher in 1991 with species found in South Africa, Australia and South America. The South African species Vibone vetusta was the only one described prior to Hunt & Cokendolpher’s (1991) paper, and until now no further species had been described since. The main reason these animals were overlooked previously is probably their size: ballarrines include some of the smallest of all harvestmen (the specimen I was looking at, for instance, has a central body only a bit over a millimetre long). Ballarrines differ from other harvestmen in the form of their pedipalps which are relatively long and have the patella much longer than the tibia (the converse is usually the case). Whereas other phalangioid harvestmen have the patella and tibia of the pedipalp more or less in a straight line or have the tibia bent slightly downwards, Hunt & Cokendolpher (1991) were struck by how the ballarrines had the tibia reflexed upwards relative to the patella. Ballarrine pedipalps also lack a terminal claw, and have only a relatively few glandular hairs instead of the denser covering of simple hairs found in other harvestmen. As noted in an earlier post and paper that I was associated with (Wolff et al. 2016), the overall pedipalp form is adapted for preying on small animals such as springtails: the long pedipalp acts like a tentacle that can be whipped forward to trap prey with its sticky hairs.
Until this point, New Zealand had been a puzzling gap in the Ballarrinae’s otherwise classic Gondwanan distribution (long-term readers may recall that this is the second time I’ve seen a puzzling biogeographical lacuna filled). I didn’t have any idea why that should be absent but even after looking at probably thousands of harvestmen specimens from all corners of the country I still hadn’t seen any. Hence my immediate excitement about the find, but said excitement was also leavened with a certain degree of caution. Harvestmen taxonomy is heavily dependent on features of the males (particularly the male genitalia) with females of closely related species often being indistinguishable. Unfortunately, the only specimen of New Zealand ballarrine I had on hand was female. Sorting through the remainder of the collection I was working on failed to turn up any more. I even considered whether I could wrangle a trip to the original collection locality to see if I could find more specimens, but that proved unfeasible. The ballarrine had been collected by J. Dugdale in 1980 at a spot called the Dart Hut, which lies at the summit of the Rees-Dart walking track in Mount Aspiring National Park. This is a pretty isolated part of the country with no permanent population and no nearby roads. Travelling to the Dart Hut by foot takes a minimum of two days each way; the usual time taken to travel the Rees-Dart is five days (its supposed to be a nice hike that travels through similar terrain to the more famous Milford Trail without the massive crowds of the latter). What is more, at the time I was looking into it, the Rees-Dart was closed until further notice due to flooding earlier in the year taking out one of the bridges along it. Nevertheless, I eventually decided that the value of publicising the presence of this significant group in New Zealand outweighed the risk of not yet being able to confirm male morphology. Unfortunately, the nature of the specimen (spindly legs everywhere!) meant that I found myself unable to get good photographs and the resulting paper had to be illustrated with (always somewhat ropey when I do them) hand-drawn illustrations; nevertheless, the best photos I got are here in this post.
Fortunately, sexual dimorphism within ballarrines tends to be low. I was very interested to see that the New Zealand ballarrine was more similar to the South American species Americovibone lanfrancoae than to any of the Australian species; so much so, in fact, that I ended up assigning it to the same genus as Americovibone remota. Americovibone lanfrancoae is also a very rare species, being described from only two known specimens from the Tierra del Fuego region. The most obvious difference between A. remota and A. lanfrancoae is that, in the former, the tibia of the pedipalp is not reflexed back above the patella as in every other ballarrine but is bent slightly downwards in a more standard position for phalangioids. This has some very interesting implications for ballarrine phylogeny. A molecular phylogenetic study of long-legged harvestmen by Groh & Giribet (2014) that included two ballarrines (the South African Vibone vetusta and the Australian Ballarra longipalpis) failed to unite the two as a clade. If accurate, this result would require the distinctive ballarrine pedipalp to have evolved on more than one occasion. The observation that A. remota may retain a more plesiomorphic pedipalp morphology could provide some correlation for this possibility.
But if Ballarrinae are indeed present in New Zealand, why are they apparently so rare? Part of the reason may be to do with habitat. Both the New Zealand and South American species of Americovibone are known from forests dominated by Nothofagus, southern beech. This tree genus is widespread in upland and colder parts of New Zealand. A bit north of the collection locality for A. remota, however, is an area where the beech forests disappear for a distance of a couple of hundred kilometres: this has been referred to as the “Nothofagus gap”. Studies on other groups of organisms show that this gap is a significant one for New Zealand biogeography, with many beech-associated species restricted to one side or the other of the gap. Could A. remota be a specialist of the south-west beech forests of the South Island? If so, it is unique to one of New Zealand’s least known corners.
Systematics of Ballarrinae
Characters (from Hunt & Cokendolpher 1991): Body small, usually greater than 0.8 and less than 3.5 mm long. Integument usually thin, at the most armed with minute setae. Abdominal spiracle usually concealed beneath coxa IV. Penis shaft usually with strong dorsal keel, muscle usually long (tendon short), left ventrolateral barbed process present or lacking. Ovipositor with two or four seminal receptacles. Pedipalp patella longer than tibia and usually tarsus, dorsal angle between patella and tibia less than 180°, patella and tibia without dense pile of setae; tarsus usually concave dorsad along its length, plumose setae extend to or almost to distal tip, non-socketed setae restricted to area of distal tip or lacking; tarsal claw lacking or small, usually inserted ventrodistally, usually with on ventral tooth. Chelicerae small in most species; first segment with or without ventral spur; jaws with continuous row of teeth of subequal size in most species. Legs long and usually slender; pseudoarticulations present in all metatarsi, tibia II, and femur II in some species; tarsi each with simple claw or claw with small teeth on each side. Sexual dimorphism slight in most species.
<==Ballarrinae |--Plesioballarra Hunt & Cokendolpher 1991HC91 | `--*P. crinis Hunt & Cokendolpher 1991HC91 |--Arrallaba Hunt & Cokendolpher 1991HC91 | `--*A. spheniscus Hunt & Cokendolpher 1991HC91 |--Americovibone Hunt & Cokendolpher 1991HC91 | |--*A. lanfrancoae Hunt & Cokendolpher 1991HC91 | `--A. remota Taylor 2016T16 `--Ballarra Hunt & Cokendolpher 1991HC91 |--*B. drosera Hunt & Cokendolpher 1991HC91 |--B. alpina Hunt & Cokendolpher 1991HC91 |--B. cantrelli Hunt & Cokendolpher 1991HC91 |--B. clancyi Hunt & Cokendolpher 1991HC91 |--B. longipalpus Hunt & Cokendolpher 1991HC91 `--B. molaris Hunt & Cokendolpher 1991HC91
*Type species of generic name indicated
References
[HC91] Hunt, G. S., & J. C. Cokendolpher. 1991. Ballarrinae, a new subfamily of harvestmen from the Southern Hemisphere (Arachnida, Opiliones, Neopilionidae). Records of the Australian Museum 43: 131–169.
[T16] Taylor, C. K. 2016. First record of a representative of Ballarrinae (Opiliones: Neopilionidae), Americovibone remota sp. nov., from New Zealand. Journal of Arachnology 44 (2): 194–198.
Wolff, J. O., A. L. Schönhofer, J. Martens, H. Wijnhoven, C. K. Taylor & S. N. Gorb. 2016. The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida, Opiliones). Zoological Journal of the Linnean Society 177 (3): 558–601.