Plates 1 and 2. Text-fig. 1. Figs. 1–7.
Macropathus filifer was first described by Walker (1869) as belonging to his new genus Macropathus. Specimens were collected from caves in New Zealand by H. Drew Esq., some being stated to occur “half a mile within”, and sent to England. Walker divides his genus into three species—M. filifer, M. fascifer and M. altus. His descriptions of M. filifer and M. fascifer are from male specimens and M. altus is from a female. From the similarity between the three descriptions, and making allowances for minor variations which have been found to occur, these three species of Walker would appear to be the male and female of the same species. As M. filifer has page priority over M. fascifer and M. altus, the name of the species must therefore become M. filifer
In New Zealand Colenso (1881) described a new species of weta from one specimen collected “in dark underground caves near the head of the Mauawatu River, in the ‘Forty mile bush’ 1879” as Hemideina speluncae. He says, “This peculiar and very interesting animal, (of which I regret to say I have but one whole specimen), inhabits in great numbers those small caves which are difficult of access; there they hop and spring about like shrimps, and having such excessively long and fine horns and legs, it is a very difficult matter to secure a perfect
specimen; of course the necessity of having a candle burning when in those dark recesses, greatly increases the difficulty.” From his description of the long slender legs of the specimen, giving the length and number of spines present on each, and the extreme length of the antennae, bearing short obtuse spines for one-third of their length it is fairly safe to say he had a male specimen of Macropathus filifer.
In 1862 Scudder erected the genus Hadenoecus and described species from Europe and North America in the Boston Journal of Natural History VII. Then in 1869 he described '' A New Cave Insect from New Zealand “in Ent. Notes Proc Bost. Soc. Nat. Hist. 12, and added it to his genus under the name Hadenoecus edwardsii. The colour and measurements Scudder gives are not sufficient to give a clear description, and he admits that it is described from “one imperfect specimen, much the largest of the genus, … presented to me by my friend Mr. Henry Edwards, who captured it himself a limestone cave at Collingwood, Massacre Bay, Middle Island, New Zealand. The cave is close to the sea shore and near a very large coal deposit, which occasionally crops out in the interior. The Hadenoeci were rather numerous but very difficult to catch, disappearing in the crevices of the rocks on the approach of lights. They appeared to be most abundant near the streams of water which percolated through the rocks. The sex of my specimen cannot be determined.”
In 1888 Brunner took H. edwardsii and made it the type species of his new genus Pachyrhamma which contained two species, P. edwardsii (Scudd. 1869) and P. novae-seelandiae n. sp. both from New Zealand. Brunner makes no reference to Walker's paper of 1869 and his genus is synonymous with Macropathus. From Brunner's descriptions P. edwardsii and P. novae-seelandiae are synonymous with Macropathus filifer and Macropathus fascifer respectively. Brunner says of P. edwardsii, “The fore femora are unarmed beneath, the posterior femora are the longest and the most graceful and beneath the inner margin are 4–6 rather long spines. The outer margin is armed with smaller spines close together.” Walker's description says, “Legs very slender, extremely long, thrice longer or more than the body; hind femora with six minute spines beneath, slender except near the base.” Both were describing males, and the similarity between the two descriptions is obvious. Of P. novae-seelandiae Brunner says, “Anterior femora armed beneath with four or five small spines. Posterior ones thickened at base, beneath, inner margin has 8–10 small spines, outer margin armed with 4 spines.” Walker says, “Legs rather slender, very long; femora and fore anterior tibiae armed with a few spines on each side beneath.” It is a pity Walker gives no record of the exact number of spines present. As Walker described Macropathus fascifer in 1869, it has priority over Brunner's new species Pachyrhamma novae-seelandiae. But Macropathus filifer and M. fascifer have been shown to be synonymous, therefore Pachyrhamma edwardsii and P. novae-seelandiae must also be synonymous. Examination of over one hundred and twenty specimens of Macropathus filifer has shown me the wide range of variability which occurs within the species, especially in the number of spines of the legs, and because of this, I have no hesitation in grouping all these species under the name Macropathus filifer Walker, 1869.
In 1952 a limestone cave near Collingwood was visited by Dr. J. T. Salmon find cave-wetas collected from it. The cave is near the sea, and it appears to agree with the description given by Scudder as the habitat of H. edwardsii. I
was allowed to examine a pair of these wetas, which were small and very difficult to catch, disappearing into the crevices of the limestone as described by Edwards. These wetas were the only species in the cave and they definitely do not belong to the genus Macropathus. The measurements of these wetas show them to be smaller than the specimen described by Scudder, but the wetas collected in 1952 may have been immature specimens. Brunner erected a new genus for the cave-weta from Collingwood, but the weta he described as his type specimen of the genus was not the one used by Scudder. Brunner describes both male and female, whereas Scudder's specimen could not be sexed. Brunner obviously had obtained specimens of the more widely distributed cave-weta Macropathus filifer. Thus Brunner's genus Pachyrhamma is a synonym of Macropathus and the wetas from Collingwood must still be called Hadenoecus edwardsii until they are placed in a new genus.
In 1897 Hutton published a paper on the Systematics of the Stenopelmatidae of New Zealand. In his introduction to this paper he says, “The cave-wetas are in the greatest confusion, and we do not know whether there are six or only two species.” He completely ignores the descriptions of Walker and Brunner and redescribes the two genera Pachyrhamma and Macropathus. He obviously used the species fascifer to redescribe the genus Pachyrhamma, but from examination of Brunner's monograph the type species of Pachyrhamma must be filifer and not edwardsii as shown by Hubbell (1936), so Hutton used the wrong specie-for his reconstruction of the genus Pachyrhamma and fascifer should belong to Macropathus.
After his description of the genus Macropathus Hutton says, “I have had to reconstruct this genus in order that it may be understood. It is very different from Pachyrhamma.” He disregards the genus Hadenoecus Scudder and places the two species Macropathus filifer Walker and Hadenoecus edwardsii Scudder in this genus. Thus it can be seen from the foregoing synonymy that Hutton crossed the names of the two genera so that Pachyrhamma should be Macropathus and Macropathus, Pachyrhamma.
With regard to the genus Pachyrhamma he says, “There is considerable confusion among the species of this genus, if, indeed, there are more than one. I shall commence with the only species that has been adequately described, and then point out the characters which may possibly separate the others from it.” The species which he describes in great detail is Pachyrhamma speluncae, the synonym of Hemideina speluncae Colenso (1881) which Colenso had put into the wrong sub-family. The other two species in his genus are P. novae-seelandiae Brunner and P. fascifer Hutton had realised that Macropathus fascifer Walker and Macropathus altus Walker were the male and female of the same species, and he was the first to call them P. fascifer (Walker). Hutton separates out his three species by differences in number of the spines on the legs. He separates P. novae-seelandiae from P. speluncae on the basis of P. novae-seelandiae having no spines on the middle femora, no spines on the upper surfaces of the fore and middle tibiae and no peculiarities in the antennae. P. novae-seelandiae is separated from P. fascifer by having four or five spines on the lower surface of the fore femora instead of two together with no mention being made of the two spines on the lower surface of the middle tibia. P. fascifer is separated from P. speluncae by no reference being made to peculiarities of the antennae and
no spines being present on the upper surface of the middle tibiae. Also the fore femora have only two spines below and the middle tibiae three in each row, while P. speluncae has five and four respectively. One thing is obvious from this, Hutton had examined very few specimens of Pachyrhamma, and he could never have collected any in the field. If he had he would have realised the great range of variability that occurs in the number of spines present on the legs of Pachyramma. So variable is this number that there often is fluctuation in the number of spines on the right and left fore, middle or hind legs. Thus it is obvious that the number of spines present could never be used as a taxonomic character. With regard to the antennae, he would have realised that the presence of spines on the proximal half of the antennae is a secondary sexual characteristic of the male and the female never possesses them. P. speluncae was described from one specimen only, a male, therefore it was natural for spines to be present on the antennae. P. novae-seelandiae was described from females only, so consequently no spines would occur. P. fascifer is described by Walker from both male aid female specimens, but it is possible the male was immature, as it is only on the antennae of fully mature male cave-wetas that spines are present. Thus Button's three species can be synonymised and as fascifer has priority because it was described first in 1869, they all become P. fascifer, which as I have already shown is correctly Macropathus filifer. Another point Hutton overlooked in his description of the genus Pachyrhamma was the presence of a short retro-lateral apical spine on the middle femora. This spine has been present on every specimen I have examined, and as the number of apical spines present on the legs of each weta is constant, it is a good generic character.
Alfken (1901) records a new member of the Rhaphidophoridae collected from Stephens Island during the Schauinsland Expedition 1896–97. The species is described from a single male and four male nymphs. No females were collected. Alfken connects this species with Hadenoecus Scudder but, because its fore and middle femora bear spines beneath and the middle femora have two apical spines, he places it in the genus Gymnoplectron Hutton, 1896. However, from the body and leg measurements he gives, the number of spines on the legs, and the particulate description of the colour pattern, it is obvious that he was describing specimens is of Macropathus filifer. This is the first record of M. filifer from Stephens Island.
In my own work on Macropathus filifer I have examined a representative number of specimens from Trio Islands, Stephens Island, Percy's Reserve, Karori and Ruakokopatuna. Tables were made from spine counts of the legs and Table I shows the wide range of variability which occurs within the species. From careful examination of this material six constant features emerged by which the species an be recognised:
Apical spines on femora and tibiae of fore and middle legs, and on femora, tibiae and two proximal segments of tarsi of hind legs are always constant.
Fore coxae only, always armed with a retrolateral spine.
Neither apical spines nor any other spines occur on fore or middle tarsi.
Subgenital plate in both male and female is always constant in shape.
Sexually mature males always bear spines on the proximal half of their antenna; females never possess them at any stage in their life history.
There is very little difference in size between males and females collected from the same locality, but there is considerable variability when specimens from different areas are compared.
The species Macropathus filifer is now redefined as follows:—
Colour. Basic colour medium to dark ochrous, with anterior and posterior borders of pronotum and posterior borders of mesonotum, metanotum and abdominal terga a dark reddish-brown; lateral and posterior margins pale ochrous; femora and tibiae of all legs transversely banded with broad bands of light brown and dark reddish-brown, interspersed with narrow bands of pale ochrous; antennae medium brown; ovipositor deep reddish-brown.
Body. Length up to 41 mm., average 35 mm. Body sparsely clothed with golden setae. Ovipositor seven-eighths as long as body. Antennae 4.5–5 times as long as body. Head vertical. Compound eyes laterad, nearly elliptical, twice as long as broad; a single anterior, white, median ocellus only. Fastigium as high as long, rising abruptly, slightly sulcate, with base touching scape of antennae. Mandibles small. Maxillary palps with third and fourth joints sub-equal in length. Pronotum rounded anteriorly and produced in front over occiput, truncated posteriorly; sternum transversely narrowed; pronotum and mesonotum distinctly margined laterally and posteriorly. Cerci Fig. 1 (C), long, tapering, unsegmented, clothed with long and short setae. Bodies of male and female from same locality subequal, but considerable variability in specimens from different localities.
Antennae. Fig. 4 Very long, slender, tapering flagellum (F), thick and almost touching at base; scape about four times as large as pedicel, which is narrower than stape, but broader than other segments; third segment on dorsal aspect narrower than pedicel, but half as long again, and on ventral aspect equal in length with pedicel; from fourth segment onwards segments unequal in length, although steadily decreasing in size; all segments thickly clothed with short golden setae. Sexual dimorphism is present in antennae, male possessing longer, stouter antennae than female; middle portion of flagellum (F) in male armed with a number of short ventral retrolateral spines (SP), each borne on a swelling.