Review of the Australian and New Zealand orb-weaving spider genus Novakiella (Araneae, Araneidae)
Author
Framenau, Volker W.
https://orcid.org/0000-0002-7724-3831
Harry Butler Institute, Murdoch University, 90 South St, Murdoch, Western Australia 6150, Australia & Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia & Centrum fuer Naturkunde (CeNak), Universitaet Hamburg, Martin-Luther-King-Platz 3, 20146, Hamburg, Germany
volker.framenau@murdoch.edu.au
Author
Vink, Cor J.
https://orcid.org/0000-0003-4237-0117
Department of Pest-management and Conservation, PO Box 85084, Lincoln University, Lincoln 7647, Christchurch, New Zealand & Centrum fuer Naturkunde (CeNak), Universitaet Hamburg, Martin-Luther-King-Platz 3, 20146, Hamburg, Germany
Author
Scharff, Nikolaj
Zoology Section, Research and Collections, Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark
Author
Baptista, Renner L. C.
Laboratorio de Diversidade de Aracnideos, Universidade do Brasil / Universidade Federal do Rio de Janeiro. Av. Carlos Chagas Filho 373, 21941 - 902, Ilha do Fundao, Rio de Janeiro, Brazil
Author
Castanheira, Pedro de S.
https://orcid.org/0000-0002-0623-1622
Harry Butler Institute, Murdoch University, 90 South St, Murdoch, Western Australia 6150, Australia & Laboratorio de Diversidade de Aracnideos, Universidade do Brasil / Universidade Federal do Rio de Janeiro. Av. Carlos Chagas Filho 373, 21941 - 902, Ilha do Fundao, Rio de Janeiro, Brazil
text
Zoosystematics and Evolution
2021
2021-07-26
97
2
393
405
http://dx.doi.org/10.3897/zse.97.67788
journal article
http://dx.doi.org/10.3897/zse.97.67788
1860-0743-2-393
A32F2CCD3CB8457087D13B38CA555E3C
564A467951EF5C8A883587B138B7809E
Genus
Novakiella Court & Forster, 1993, in Platnick (1993)
Novakia
Court & Forster, 1988: junior homonym of
Novakia
Strobl, 1893 (
Diptera
) and
Novakia
Tolmachoff, 1926 (
Mollusca
) is another junior homonym.
Type-species.
Epeira tri-tuberculata
Urquhart, 1887.
Diagnosis.
The informal clade of the backobourkiines is well supported by the molecular phylogeny of
Scharff et al. (2020)
, but the taxonomy and systematics of the species and genera within this clade are poorly resolved. Only three genera within the clade have been revised using modern taxonomic methods:
Plebs
,
Backobourkia
and
Lariniophora
Framenau, 2011. The genera
Carepalxis
and
Acroaspis
have not been revised and their putative synapomorphies remain unknown. It is therefore difficult to diagnose
Novakiella
against these genera. Other Australian backobourkiines included in
Scharff et al. (2020)
represent species that have clearly been misplaced in genera they do not belong to (i.e.,
Eriophora
or
Araneus
) and these represent undescribed genera (in that study listed as
"NGEN01"
for
Eriophora transmarina
(Keyserling, 1865),
"NGEN02"
for
Araneus recherchensis
Main, 1954 and
"NGEN05"
for
Araneus senicaudatus
Simon, 1908). Until these species have been revised and placed in new or existing genera,
Novakiella
cannot be diagnosed against them.
Novakiella
distinctly differs from the revised backobourkiine genera by overall somatic morphology. The abdomen is subtriangular with strong humeral humps (Figs
1A
,
3A
,
4A
,
6A
), while it is rounded with small humerals in
Backobourkia
(i.e.,
Framenau et al. 2010
, fig. 5), slightly elongated in
Plebs
(e.g.
Joseph and Framenau 2012
, figs 6, 7, 10), and strongly elongated in
Lariniophora
(
Framenau 2011
, figs 2, 3). Males can be differentiated by the presence of a tibial apico-prolateral spur carrying a thick spine (or macroseta) on leg II (Figs
1E
,
4C
).
Verrucosa
McCook, 1888 and
Carepalxis
also have a spur on leg II, but in both genera it carries two spines (
Levi 2002
: p. 546;
Lise et al. 2015
: p. 5; VWF pers. obs.). In addition,
Verrucosa
is limited to the Neotropics and not part of the backobourkiines (
Scharff et al. 2020
). There are distinct differences in the male pedipalp morphology between
Novakiella
and other backobourkiines.
Novakiella
males have a stout median apophysis that is drawn out into a basally pointing acute projection (Figs
1C
,
2B
,
4D, E
,
5
); in contrast, the median apophysis has a basal flange in
Backobourkia
(
Framenau et al. 2010
) (absent in
Novakiella
), is elongate transverse with two apical tips in
Plebs
(Joseph & Framenau, 2012, e.g. figs 4B, 8A), and has a two-humped lobe in
Lariniophora
(Framenau, 2011, fig. 4). All backobourkiines appear to have a basal extension of the conductor (discussed in
Framenau et al. 2010
and there termed paramedian apophysis), but in
Novakiella
it is very different to all other described backobourkiines and much more conspicuous; we here propose a new term, conductor lobe (CL), which extends apically well past the radix (Figs
1C
,
2A, B
,
4D-F
,
5
).
Figure 1.
Novakiella trituberculosa
, male (WAM T73571).
A.
Dorsal habitus;
B.
ventral habitus;
C.
Left pedipalp, mesal view;
D.
Left pedipalp, ventral view. Abbreviations: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; P, paracymbium; Ra, radix; TA, terminal apophysis. Scale bars:
A, B
, 1 mm;
C, D
, 0.5 mm;
E
, 1.2 mm.
Figure 2.
Novakiella trituberculosa
, male (WAM T73571), expanded left pedipalp.
A.
Ventral view;
B.
Retrolateral view. Abbreviations: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; P, paracymbium; Ra, radix; TA, terminal apophysis. Scale bars: 0.5 mm
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
3C
,
6C
;
Framenau et al. 2010
, e.g. figs 6D, F;
Framenau 2011
, fig. 6;
Joseph and Framenau 2012
, e.g. figs 4D, 8E).
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
1A
,
3A
,
4A
,
6A
); fovea longer than wide in males and wider than long in females, and with a dark spot in both sexes (Figs
1A
,
3A
,
4A
,
6A
); colouration (of ethanol preserved specimens) varying from reddish-brown to yellowish-brown, with black patches along carapace borders (Figs
1A
,
3A
,
4A
,
6A
). Eyes ringed in black, anterior median eyes largest, posterior eye row slightly recurved, lateral eyes almost touching, posterior lateral eyes separated from posterior median eyes by more than their diameter and located on small tubercles at the clypeus border (Figs
1A
,
3A
,
4A
,
6A
). Chelicera paturon with dark hue, fangs reddish-brown. Labium wider than long, subtriangular, with front end bulging and beige (Figs
1B
,
3B
,
4B
,
6B
). Endites rounded, inner portion beige (Figs
1B
,
3B
,
4B
,
6B
). Sternum almost as long as wide with dark contour (Figs
1B
,
3B
,
4B
,
6B
). Legs (Figs
1A, B, E
,
3A, B
,
4A-C
,
6A, B
): Leg formula IV> I> II> III, all longer than
body's
length with dark spots on joints; tibia II of males with apico-prolateral spur bearing a thick macroseta or spine (less pronounced in
N. boletus
sp. nov.). Abdomen subtriangular, longer than wide, with two distinct humeral humps and posterior tip reaching beyond spinnerets (Figs
1A
,
3A
,
4A
,
6A
); folium pattern distinct; sides varying in colour from yellowish-brown to black (Figs
1A
,
3A
,
4A
,
6A
), venter light coloured, generally mottled dark (Figs
1B
,
3B
,
4B
,
6B
). Male genitalia (Figs
1C, D
,
2A, B
,
4D-F
,
5
): male pedipalp patella with a single strong macroseta; paracymbium well-developed and hook-like; cymbium longer than wide; radix thick and elongated, reaching from the base of median apophysis to near the cymbium tip; conductor lobe conspicuous and projected apically, being composed of two distinct lobes (
N. trituberculosa
) or mushroom-shaped (
N. boletus
sp. nov.); terminal apophysis wider than long, rounded and tapering terminally; conductor well-developed, subquadrate; embolus uncapped, elongated, pointed and almost straight; median apophysis stout, with an acute basally pointing tip. Female genitalia (Figs
3C-E
,
6C
): epigyne plate wider than long, subtriangular; scape much longer than wide and extending posteriorly beyond plate (but length not known in
N. boletus
sp. nov.), generally broken off. Spermathecae spherical and occupying most of genital area.
Figure 3.
Novakiella trituberculosa
, female (WAM T73556).
A.
Dorsal habitus;
B.
Ventral habitus;
C.
Epigyne, ventral view;
D.
Cleared epigyne, ventral view;
E.
Cleared epigyne, dorsal view. Abbreviations: CO, copulatory openings; S, scape; Sp, spermatheca. Scale bars:
A, B
, 2 mm;
C-E
, 0.1 mm.
Composition.
Novakiella trituberculosa
(Roewer, 1942) and
N. boletus
sp. nov.
Figure 4.
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.
Remarks.
The nomenclatural history of
Novakiella
is convoluted.
Novakiella trituberculosa
was first described as
Epeira tri-tuberculata
by
Urquhart (1887)
, before
Roewer (1942)
replaced the species-group name as it is a junior primary homonym of
Epeira trituberculata
Lucas, 1846, currently listed as a junior synonym of
Cyclosa insulana
(Costa, 1834).
Court and Forster (1988)
described the genus
Novakia
to accommodate this species; however, this new genus-group name was also preoccupied, by
Novakia
Strobl, 1893 (
Diptera
) and
Novakia
Tolmachoff, 1926 (
Mollusca
). Court and Forster, in
Platnick (1993)
, proposed
Novakiella
Court & Forster, 1993 as a replacement name.
Figure 5.
Novakiella boletus
sp. nov., male holotype (MV K9867), expanded left pedipalp. mesal view. Abbreviations: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; Ra, radix; TA, terminal apophysis. Scale bar: 0.5 mm.
Figure 6.
Novakiella boletus
sp. nov., female (SAM).
A.
Dorsal habitus;
B.
Ventral habitus;
C.
Epigyne, ventral view. Abbreviations: CO, copulatory openings; L, lips; S, scape. Scale bars:
A, B
, 2 mm,
C
, 0.2 mm.
Distribution.
Australia and New Zealand (Figs
7
,
8
).
Figure 7.
Distribution records of
Novakiella trituberculosa
(green circles) and
Novakiella boletus
sp. nov. (red stars) in Australia.
Figure 8.
Distribution records of
Novakiella trituberculosa
in New Zealand; red circle points to material examined for this study and yellow diamonds point to records literature records (
Court and Forster 1988
).