Research Article |
Corresponding author: Renzo Perissinotto ( renzo.perissinotto@nmmu.ac.za ) Academic editor: Steven Lingafelter
© 2016 Anders Bjørnstad, Elizabeth Grobbelaar, Renzo Perissinotto.
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:
Bjørnstad A, Grobbelaar E, Perissinotto R (2016) Review of Afraustraloderes rassei Bouyer, 2012: description of its female and a new species of Pixodarus Fairmaire, 1887 (Coleoptera, Cerambycidae, Prioninae). ZooKeys 558: 77-93. https://doi.org/10.3897/zookeys.558.6112
|
The original description of Afraustraloderes rassei Bouyer, 2012 included a female that is now recognized as a separate species belonging to the genus Pixodarus and here described as P. spiniscapus sp. n. The true female of A. rassei has also been obtained recently and is, therefore, here described. The synonymy of P. exasperatus with P. nyassae, proposed earlier by
Hopliderini , new species, Cape Floral Region, South Africa
Afraustraloderes rassei Bouyer, 2012 was originally described based on a small collection of three males and one female from South Africa.
The male and female specimens represented in the photos of Figures
In 2006, one of the authors (EG) was working on the collection of the late Richard Watmough, a well-known South African entomologist who passed away in 2005 (
Adult specimens of A. rassei were obtained either through direct searching of burnt and dead logs/roots of Proteaceae in the fynbos vegetation or by rearing their larvae in a controlled, closed environmental room in Port Elizabeth, about 170 km from the Joubertina collection locality. Larvae were collected in an advanced stage of development and maintained in the logs in which they were found in closed plastic crates, in darkness and at ambient temperature. Logs were left lying in a horizontal position with minimal manipulation and moistened by spraying rainwater on the outer surface once a month.
Specimen length was measured from the anterior margin of the head to the elytral apex and the width was measured at the widest point of the elytra. Photos of set specimens were taken using either a Canon EOS 5D camera fitted with a Canon MP-E 65 Macro 2.8–1.5× objective or a Canon PowerShot G11 with automatic macro setting. The background was removed from the photos using Microsoft Word 2010 (Picture Tools), to increase clarity of resolution. Combine ZP Image Stacking Software by Alan Hadley (alan@micropics.org.uk) was used to obtain z-stacking composite images. In situ photos were taken using a Ricoh CX1 camera with macro setting.
Terminology used to describe the male genitalia follows the standard work of
Holotype label data is quoted verbatim. Collections are abbreviated as follows:
Four observations of the new Pixodarus species were located on the citizen science platform iSpot (
Type species. Afraustraloderes rassei Bouyer, 2012
When
The tribe Hopliderini was revised by
eyes strongly emarginate, but not embracing antennal tubercle;
eyes widely separated dorsally;
antennae as long as or exceeding the body length in males;
antennae gradually becoming more slender distally;
scape very short;
3rd antennomere very long;
anterior coxal cavities open posteriorly;
prosternal process pointed;
metathoracic episterna usually truncate apically, without posterior constriction;
tibiae apically spined;
pronotal disc distinctly punctate;
lateral margins of pronotum each bearing five spines;
anterior border of pronotum with a silky tuft on each side of the ‘neck’;
head strongly declivous, resulting in mandibles not being visible in dorsal view.
Afraustraloderes has some of these characteristics: viz. items 1, 2, 5, 7, 13 (partly) and 14. But it does not exhibit the majority of the features listed above: viz. nos. 3, 4, 6, 8, 9, 10, 11, 12. Regarding no. 9, although the metathoracic episterna are truncate apically, there is an evident constriction near the distal end. Regarding no. 10, there are the usual two spurs between the end of the tibia and the base of the tarsus, but there are no apical teeth or spines on the tibiae themselves. Regarding no. 11, there is micro-punctation on the disc of the male, but not in the female. Thus, from the above analysis it can be concluded that Afraustraloderes does not belong in the tribe Hopliderini. We propose that this genus be removed from the Hopliderini and regarded as Prioninaeincertae sedis.
Afraustraloderes rassei Bouyer, 2012: 214
Paratypes: 1♂ RSA, ECA, Joubertina (33°53'10” S, 23°50'18” E), 18 Dec 2009, M Villet and R Smith leg (RPPC); 1♂, same locality, but 25 Dec 2010, R Perissinotto and L Clennell (TBPC). Other material: 1♂, same locality as paratype, but Oct 2010, found dead on the ground, Rodger Smith leg (RPPC); 1♀, same locality, but 28 Dec 2014, found dead inside dead Protea log, R Perissinotto and L Clennell (RPPC); 5♂ 1♀, same locality, bur reared from larvae in Port Elizabeth, adults emerged 23 Dec 2014-7 Jan 2015 (ABPC, NDPC,
Size. 29.6–31.5 mm long including pygidium, 11.5–12.2 mm wide (maximum width at metacoxae).
Head. Mandibles relatively short and broad with foveolate base, distal part short, shiny and weakly arcuate with pointed apex, cutting edge with a small tooth near base; maxillary and labial palpi with terminal segments terete and slightly truncate at apex; galeae much shorter than palpi and covered in rufous bristles; clypeus with long stiff rusty brown bristles directed anteriorly; frons very uneven; antennal tubercles moderately raised, with strongly uneven, deeply sculptured surface; eyes small, finely facetted with deep emargination dividing each eye into subequal lobes, with lower lobe almost reaching gula; vertex strongly uneven and deeply sculpted, with poorly defined median depression.
Antenna. Short and slender, only reaching slightly beyond humeri of elytra; with scape as longest antennomere, finely punctate with a narrow base gradually widening distally and ending in apical spine on posterior margin; antennomeres 2–10 with circular cross-section, exhibiting narrow base but widened apically; 11th antennomere slightly compressed; pedicel very short, 3rd antennomere almost as long as scape, antennomeres 4–11 subequal in length.
Pronotum. Distinctly transverse, with disc deeply sculpted by irregular reticulations and foveolations; two strongly uneven, raised areas with smooth and shiny surface present about halfway between anterior and posterior margin; anterior margin with fringe of short, black setae curled around base of head, but slightly longer and straight on either side of head, without forming a marked ‘brush’; lateral margins with many (15–20) short teeth, foremost tooth much stronger than others and directed forwards, last tooth on each side, near posterior corner, much larger and sharper than the rest.
Scutellum. Broadly tongue-shaped to triangular, with finely reticulate surface.
Elytron. Shallowly reticulate basally, sculpture becoming less distinct towards rounded apices; humeri smoothly rounded.
Legs. Short and slender; femora only slightly thickened at middle, hardly projecting beyond lateral borders of elytra; tibiae nearly straight, only slightly curved in basal part; tarsi with first tarsomere slightly longer than others, noticeably so in metatarsi.
Pygidium. Tergite 8 long, protruding beyond elytral apices.
Ventral surface. Base of maxilla punctate and shiny; submentum transversely ridged; gula glabrous, deeply punctate/reticulate/foveolate, but posterior part raised, exhibiting black bristles and less sculpture; prosternum shiny, transversally strongly convex, punctate, glabrous and with ligulate prosternal process bent dorsally at apex; proepimeron only weakly punctate, glabrous, not reaching prosternal process (i.e. procoxal cavities open); mesosternum punctate and with longitudinal median furrow, with soft black pubescence, exhibiting mesial depression just in front of mesocoxae; mesosternal process short, strongly excavate, resulting in bifurcate apex; mesocoxae moderately raised; metasternum transversally strongly convex, with median furrow increasing in depth posteriorly; whole metasternum and mesepisterna covered by soft, sparse blackish pubescence; metathoracic episterna posteriorly narrowed and truncate; all five visible abdominal sternites subequal in length, densely, but shallowly punctate, glabrous in median part, increasingly pubescent laterally with very short and soft whitish bristles; last visible sternite with evenly rounded posterior margin.
Ovipositor. Abdominal segments 8 + 9 (Figure
Sexual dimorphism. The morphological differences between the sexes are largely confined to the anterior parts of the thoracic segments. The most noticeable difference lies in the appearance and structure of the pronotum. Although the outline is quite similar in both male and female. The pronotal disc of the male has an even, matte surface, while that of the female has a very irregular lustrous surface (Figure
Ventrally the male prosternum and proepimera have a relatively smooth surface, with only a shallow microstructure giving a matte appearance (Figure
As usual, the visible abdominal sternites are transversally more convex in the female than in the male (Figures
Male genitalia.
Hoplideres nyassae Bates, 1878
In their revision of the African and Madagascan Hopliderini (then referred to as subtribe Hopliderina),
In spite of these cautionary remarks,
Pixodarus nyassae is known from RSA and Mozambique northwards to Zimbabwe and Malawi, while P. exasperatus occurs in Tanzania, southern Kenya and the eastern part of the Democratic Republic of Congo (
On the other hand, the female erroneously included in the description of A. rassei by
Holotype ♂: SOUTH AFRICA: GAU, Florauna, Pretoria 25°41'20"S, 28°09'30"E, XII.1985, R.H. Watmough (
1♂: A69 [no further data] (
1♂: RSA, GAU, Roodepoort, Walter Sisulu National Botanical Garden (26°05'21"S, 27°50'40"E), 05 Nov 2012, A Hankey, http://www.webcitation.org/6ZkmxK0on; 1♀: RSA, GAU, Randburg, Curro Aurora Private School, school grounds (26°04'48"S, 27°56'06"E), 30 Oct 2014, A Hankey, http://www.webcitation.org/6ZknFLtcm; 1♂: RSA, MPU, Presidentsrus (25°45'33"S, 29°19'05"E), 31 Oct 2014, R Bate, light trap, http://www.webcitation.org/6ZknAyhMq; 1♀, ditto, but 14 Nov 2014, http://www.webcitation.org/6ZknGm829.
Size. Male: 31.5–33.7 mm long, 12.5–13.4 mm wide (maximum width at metacoxae); female: 33.9-34.0 mm long, 13.0-13.7 mm wide (maximum width at metacoxae).
Head. Mandibles black, short and stout with deeply punctate base and strongly hooked apex; maxillary palpi tetramerous, labial palpi trimerous, basal segment of maxillary palpi extremely short and appearing 3-segmented; shape and size of the two types of palpi very similar, both shiny and brown with long, stiff, yellow setae, terminal segments hyaline in apical third and abruptly truncate; frons short, deeply concave, bordered by lateral ridges formed as continuation of mandibular bases; antennal tubercles prominent, separated by a narrow depression which continues well past the eyes posteriorly, then disappears gradually on vertex; yellowish to orange arcuate to curled bristles present on most surface areas, particularly well-developed along above mentioned depression; entire fronto-dorsal surface heavily sculpted with irregular irrorations and foveolations; eyes finely facetted, sinuately emarginate resulting in smaller dorsal eye lobe and larger suborbicular ventral lobe, emargination itself densely pubescent; genae very short.
Antenna. Scape almost wedge-shaped, gradually widening distally from narrow base, slightly compressed, with marked apical spine on proximal side; pedicel very short (< 1/5 length of scape); antennomere 4 noticeably shorter than 3 and 5 in male, not in female; antennomeres 3 and 5–10 of subequal length in male; last antennomere longest by far, more than 1.5 times length of preceding; antennomeres 3–9 + 11 of subequal length in female, only antennomere 10 slightly shorter; antennomeres 1–3 punctate in both male and female, segment 4 transitional and last 7 antennomeres only very finely micropunctate; antennomeres 2–4 practically terete, from segment 5 onwards increasingly flattened and with weak lateral tooth apically; apical segment sharply carinate; antenna reaching approximately 4/5 of elytral length or slightly longer in male, only about ½ or slightly longer in female.
Pronotum. Distinctly transverse, length/width ratio of pronotal disc (lateral spines excluded) approximately 0.6; anterior margin straight to shallowly concave, posterior margin slightly convex medially; three stout spines present laterally on either side, two straight spines at edge of anterior margin and at posterior corner respectively, one strongly curved spine between previous two and pointing posteriorly; few irregular small teeth between first and second spine; pronotal disc irregularly folded and foveate to shallowly vermiculate, elevated parts shiny; sparse yellowish- to rusty brown and somewhat curly pubescence covering much of surface, particularly dense along anterior margin.
Scutellum. Shield-like, pubescent and finely punctate.
Elytron. Dark brown, and rather elongate, ratio elytral length:combined elytral width at metacoxae varying between 1.9 and 2.3; surface distinctly costate, with three costae normally visible at least in part; humeri rounded and weakly marked; posthumeral lateral margin bent outwards and upwards, forming a miniature “gutter”; posterior sutural corner sharply angled, or exhibiting small tooth, otherwise lateral margin evenly rounded; shallow vermiculations in subscutellar part, gradually becoming finely punctate laterally and posteriorly; basal parts sparsely covered with short yellowish pubescence, gradually becoming less conspicuous posteriorly; greatest elytral width in female about half way down the elytron, distinctly wider than in male.
Legs. Profemora with a somewhat scabrid dorsal surface, ventral face punctate; meso- and metafemora smooth and lustrous dorsally; tibiae straight with short rusty brown pubescence, all widening towards the apex, with two short spurs on proximal side and blunt tooth on distal side; all tarsomeres on all legs of subequal length.
Ventral surface. Gula strongly concave and densely punctate in anterior part, transversely undulate in posterior part; prosternum pubescent, distinctly convex with anterior border strongly thickened; prosternal process ligulate with apex bent dorsally; procoxal cavities open; mesosternum punctate and pubescent, very short; mesosternal process short with median furrow, bifurcate apically; metasternum strongly convex, pubescent, finely punctate and glossy; all five visible abdominal sternites of subequal length, finely punctate, pubescent and glossy; last visible sternite with rounded to weakly truncate posterior border in female, weakly concave in male; epipleura well developed in correspondence with dorsal gutter-like extension of elytral margin.
Male genitalia. Median lobe (sensu
The name ‘spiniscapus’ refers to the prominent apical spine present on the first antennomere or scape.
Pixodarus spiniscapus sp. n. differs from P. nyassae (Bates, 1878) (syn. P. exasperatus Quentin and Villiers, 1972) in several diagnostic characters. Unlike P. nyassae, P. spiniscapus has costate elytra and these are also more elongate than in P. nyassae. Its ratio of elytral length: combined width at humeri varies from 1.9 to 2.3, while it ranges between 1.7 and 1.9 in P. nyassae. The first antennomere, in particular, has a distinct apical spine in P. spiniscapus, while it is unarmed in P. nyassae. Furthermore, in P. spiniscapus most of the pronotal disc and elytral surfaces have a yellowish pubescence, which is lacking in P. nyassae. The pronotum in P. spiniscapus has three lateral spines on either side, while there are usually five in P. nyassae. There are also significant genitalic diagnostics. While the median lobe of P. spiniscapus is very similar to that of P. nyassae, the ventral edge of the median orifice (ventral plate) in particular has an acuminate apex longer than that of P. nyassae. In the terminology of
This study shows that the two sexes originally described as belonging to the same species, A. rassei, actually represent two very different species. The unique characteristics of the genus Afraustraloderes are also highlighted, revealing the unlikelihood of it belonging to the tribe Hopliderini. According to the analysis undertaken by
The habitat and ecology of A. rassei are also unusual for a prionine – both adults and larvae have so far only been found in fynbos vegetation (Figure
Among the fynbos vegetation, adults and larvae of A. rassei were found on or inside trunks, branches and upper roots of dead Protea and Leucadendron species (Proteaceae) and also Hakea sericea Schrad. and J.C.Wendl., which is an alien invasive plant in the same family, introduced into South Africa from southeastern Australia in the 19th century (
Larval stages of A. rassei were first collected in December 2010, but were small, < 15 mm long, and did not survive to the next season in captivity. By December 2013, larvae observed in their natural habitat had reached an average size of 30–50 mm and were able to survive in captivity, with a few eventually pupating and emerging as adults in December 2014–January 2015. Others are currently completing their life cycle in the laboratory. As the fynbos area where larvae were monitored had been burned by a veld fire in the winter of 2009, it would appear that larvae may take about 6–7 years to complete their development to imago. Thus, it seems likely that this reflects the cycle of natural fire burning in the fynbos vegetation of this region (
Unfortunately, very little is known about the habitat and biology of Pixodarus spiniscapus, as all known specimens are accompanied by essential data only. It is most likely that the specimens collected by RH Watmough in the Florauna suburb of Pretoria were attracted to the light at night, as this is the typical behavior shown by the other known species of the genus Pixodarus (RP, pers. observ.), as also confirmed by the iSpot observations. The predominant vegetation of the Magaliesberg slopes, where the bulk of the type specimens of P. spiniscapus were collected, is of the Gold Reef Mountain Bushveld type (
We wish to thank the Eastern Cape Department of Economic Affairs, Environmental Affairs and Tourism for granting collection permits (CRO 125/14CR) and Rodger Smith, Natasja Wortel and Derek Clark for providing expert assistance and logistical support during the fieldwork undertaken in the Joubertina area. Our sincere thanks are due to Karsten Sund of the Natural History Museum, University of Oslo, Norway, and Lynette Clennell of Port Elizabeth, South Africa, for taking the photographs used in the figures of the manuscript. We thank ARC-PPRI and the Department of Science and Technology, South Africa, for funding and supporting the study, including the publication of this manuscript. Thanks are also due to Riaan Stals (ARC-PPRI) for alerting us to the iSpot records and constructive suggestions on the manuscript.