Research Article |
Corresponding author: Volker W. Framenau ( volker.framenau@murdoch.edu.au ) Academic editor: Danilo Harms
© 2021 Volker W. Framenau, Cor J. Vink, Nikolaj Scharff, Renner L. C. Baptista, Pedro de S. Castanheira.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Framenau VW, Vink CJ, Scharff N, Baptista RLC, Castanheira PS (2021) Review of the Australian and New Zealand orb-weaving spider genus Novakiella (Araneae, Araneidae). Zoosystematics and Evolution 97(2): 393-405. https://doi.org/10.3897/zse.97.67788
|
The orb-weaving spider genus Novakiella Court & Forster, 1993 (family Araneidae Clerck, 1757) is reviewed to include two species, N. trituberculosa (Roewer, 1942) (type species, Australia and New Zealand) and N. boletus sp. nov. (Australia). Novakiella belongs to the informal, largely Australian ‘backobourkiine’ clade and shares with the other genera of the clade a single macroseta on the male pedipalp patella and a median apophysis of the male pedipalp that forms an arch over the radix. The proposed genus synapomorphies are the presence of a large basal conductor lobe expanding apically over the radix and the shape of the median apophysis, which extends into a basally directed, pointy projection. Males have an apico-prolateral spur on the tibia of the second leg that carries a distinct spine. Females have an epigyne with triangular base plate bearing transverse ridges and an elongate triangular scape, which is almost always broken off. The humeral humps of the abdomen are distinct. Novakiella trituberculosa build characteristic dome-shaped webs; however, the foraging behaviour and web-shape of N. boletus sp. nov., currently only known from museum specimens, are not known.
dome-shaped orb-web, new species, systematics, taxonomy
A recent multi-gene molecular phylogenetic analysis of the orb-weaving spider family Araneidae Clerck, 1757 recovered a well-supported Australasian clade informally termed ‘backobourkiines’ (
Morphologically, the backobourkiines are still poorly circumscribed, although a single macroseta on the male pedipalp patella and the median apophysis forming an arch over the radix seem to morphologically unite all genera against many clades previously considered part of the traditional Araneinae (
The orb-weaving spider genus Novakiella was not diagnosed in detail when initially described (sub Novakia Court & Forster, 1988). However, the original description of the genus pointed to a peculiar genital morphology and an unusual horizontal orb-web “drawn up into a cone by threads attached to the hub” built by the type species, N. trituberculosa (Roewer, 1942) (
Descriptions and terminology follow recent publications on Australian orb-weaving spiders (e.g.,
Images of specimens were taken in different focal planes with a Nikon D300 digital SLR camera attached via a C-mount adapter from LM-Scope (http://www.lmscope.com) to a Leica M16A stereomicroscope and combined with Auto Montage (vers. 5.02) software from Syncroscopy to increase depth of field. We used 2 Nikon R1C1 wireless speedlights instead of fibre optics to illuminate the exposures. The latter were used as guide-light for focusing. Microscopic images of cleared epigynes were taken in different focal planes (ca. 20–30 images) on a Leica DMC4500 digital camera mounted to a Leica M205C stereomicroscope and combined using the Leica Application Suite X, v. 3.6.0.20104. Images of expanded pedipalps of N. trituberculosa were taken with a BK Plus Laboratory System from Visionary Digital (Palmyra, PA, USA) equipped with a Canon EOS 7D camera. All photos were edited and mounted with Photoshop CC 2020.
Maps were compiled in the software package QGis v. 2.14.0 Girona (https://qgis.org/en/site/; accessed 21 January 2020). Geographic coordinates were extracted directly from original labels or the registration data as provided by the museums. When no detailed geographic information was available, localities were estimated based on Google Earth v. 9.1.39.3 (https://earth.google.com/web/; accessed 21 January 2021).
Collections
MV Museums Victoria, Melbourne, Australia;
SAM South Australian Museum, Adelaide, Australia;
Morphology
Males: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; P, paracymbium; Ra, radix; TA, terminal apophysis. Females: CO, copulatory openings; S, scape; Sp, spermatheca.
Order Araneae Clerck, 1757
Family Araneidae Clerck, 1757
Novakia Court & Forster, 1988: junior homonym of Novakia Strobl, 1893 (Diptera) and Novakia Tolmachoff, 1926 (Mollusca) is another junior homonym.
Epeira tri-tuberculata Urquhart, 1887.
The informal clade of the backobourkiines is well supported by the molecular phylogeny of
Novakiella distinctly differs from the revised backobourkiine genera by overall somatic morphology. The abdomen is subtriangular with strong humeral humps (Figs
Novakiella trituberculosa, male (
Females of Novakiella have an elongated triangular scape without terminal pockets, as is typical for all backobourkiines above; however, these genera lack the subtriangular base plate with its transverse and lateral wrinkles (Figs
Description. Medium-sized (TL males ca. 5–9, females 8–12) orb-weaving spiders with males on average slightly smaller than females. Carapace longer than wide, pear-shaped; cephalic area similar in shape in both sexes (Figs
Novakiella trituberculosa (Roewer, 1942) and N. boletus sp. nov.
Novakiella boletus sp. nov., male holotype (MV K9867). A. Dorsal habitus; B. Ventral habitus; C. Left tibia, ventral view; D. Left pedipalp, mesal view; E. Left pedipalp, ventral view, F. left pedipalp, apical view. Abbreviations: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; P, paracymbium; Ra, radix; TA, terminal apophysis. Scale bars: A, B, 2 mm; C, 1 mm; D–F, 0.5 mm.
The nomenclatural history of Novakiella is convoluted. Novakiella trituberculosa was first described as Epeira tri-tuberculata by
Australia and New Zealand (Figs
Epeira tri-tuberculata
Epeira tri-tuberculata
Aranea trituberculosa
Novakia trituberculata
Novakiella trituberculosa
Court and Forster, in
Syntypes of Epeira tri-tuberculata Urquhart, 1887, 2 females, Karaka, New Zealand (37°06'S, 174°53'E), A.T. Urquhart (
AUSTRALIA: Australian Capital Territory: 1 female, 1 juvenile, Canberra, 35°18'S, 149°08'E (SAM); 1 female, Kaleen, Canberra, 35°17'S, 149°13'E, 25.iv.1990 (SAM 31192); 1 male, 1 female, Kaleen, Canberra, 35°17'S, 149°13'E, 21.v.1988 (SAM); 1 male, Red Hill, 14 Pera Place, 35°20'S, 149°08'E, 24.ii.1982, M. S. Harvey leg. (
Male N. trituberculosa can easily be distinguished from N. boletus sp. nov. by the much stronger apico-prolateral spur on tibia of leg II and the morphology of key pedipalp sclerites (Figs
Male (
Female (
The study was conducted over many years and at different institutions and therefore imaging and descriptive work based on variable specimens (plural) availability at the time. This explains why the male N. trituberculosa is here redescribed based on two specimens; one imaged many years ago, but not measured, and the measurements added for a second specimen more recently.
In Australia, mature males of N. trituberculosa were found between November and June, with peaks in January and April. Mature females were found all year round with the lowest numbers of records in November and December. Here, the species is mainly found in “pastoral habits” and constructs a horizontal orb-web amongst low grasses or weeds, with the centre pulled up by stabilizing threads. The webs are up to 0.1 m above ground. Additionally, habitat descriptions on specimen labels include “woodland”, “open forest”, “shrubs near ground”, “in long grass”, but the species also seems to occur in more disturbed habitats such as “among garden rubbish”, “ex toilet”, “walking on wall at night”, “inside house on wall”, “stationary on door knob”. In New Zealand it is mostly found in pastoral habitats (
Novakiella trituberculosa has been recorded from all Australian states, except Northern Territory, south of ca. 22°S Latitude (Fig.
Holotype male from Maits Rest, 10 km W of Apollo Bay, Otway Ranges, Victoria, AUSTRALIA, 38°45'S, 143°34'E, 16.iii.1992, G. Milledge leg. (MV K9867).
AUSTRALIA: New South Wales: 1 male, Coolah Tops National Park, off Gemini Road Loop, 31°48'59"S, 150°10'31"E, beating, 12–13.iv.2010, M. G. Rix & D. Harms leg. (
The specific epithet is a Latin noun in apposition – boletus – meaning mushroom and it refers to the distinctly mushroom-shaped conductor lobe that is reminiscent of a chanterelle (Cantharellus spp.).
Male N. boletus sp. nov. can be distinguished from N. trituberculosa by the weaker apico-prolateral spur on the tibia of leg II (Fig.
Male (based on holotype, MV K9867): Total length: 6.44. Carapace (Fig.
Female (SAM; from Kelly Hill Caves camping area, Kangaroo Island): Total length 8.5. Carapace (Fig.
Mature males of N. boletus sp. nov. were collected between February and April, females were found in May and November. Habitat descriptions include Nothofagus cunninghamii (Myrtle Beech) forest, Eucalyptus amygdalina coastal forest, and “eucalypt forest with tree fern gully”, suggesting this species occurs predominantly in temperate forests and rainforests.
Males and female N. boletus sp. nov. have not been found together, but somatic features such as size range, carapace (Fig.
This new species is only known from Australia, specifically New South Wales, South Australia, Victoria and Tasmania (Fig.
Araneidae are a highly diverse family with generally complex male pedipalp morphology which is traditionally used to infer phylogenetic relationships within this family as well as in other spiders (
In Novakiella, a basal conductor lobe is also present, but it is shaped very differently to that in other backobourkiines and the homology of a variety of structures in araneids in such a position remains unclear (
Novakiella trituberculosa was originally described from New Zealand, but
We acknowledge the support of all museum curators and scientists who facilitated loans of specimens or visits to their respective institutions: Graham Milledge and Helen Smith (