Research Article |
Corresponding author: Benjamin Wipfler ( benjamin.wipfler@uni-jena.de ) Corresponding author: Reinhard Predel ( reinhard.predel@uni-koeln.de ) Academic editor: Ben Price
© 2018 Benjamin Wipfler, Tobias Theska, Reinhard Predel.
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:
Wipfler B, Theska T, Predel R (2018) Mantophasmatodea from the Richtersveld in South Africa with description of two new genera and species. ZooKeys 746: 137-160. https://doi.org/10.3897/zookeys.746.14885
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Two new species and two new genera (Kuboesphasma, Minutophasma) of Mantophasmatodea that occur in the Richtersveld region of South Africa are described. Kuboesphasma compactum gen. n., sp. n. was found only in a small area near the village of Kuboes, while Minutophasma richtersveldense gen. n., sp. n. apparently inhabits a larger area in the Richtersveld. With these two new species, a total of four different mantophasmatodeans are known to live in this area. This is a remarkable exception to the remaining representatives of this order, where even a common occurrence of only two species is rare. We discuss this sympatry in the context of the phylogeny of the group. Additionally, we provide a map of the known distributions and a table with the most important taxonomic features of the mantophasmatodeans in the Richtersveld.
heelwalkers, Polyneoptera , lower neoptera, Kuboesphasma , Minutophasma , taxonomy, South Africa
Mantophasmatodea were newly described in 2002 (
So far 18 species of Mantophasmatodea have been described (
Here we describe the two new genera and two new species of Mantophasmatodea from the Richtersveld and present detailed information on how to distinguish the four sympatric species in this area.
The used terminology follows
The information for the specimens is given in a standard manner, i.e., locality, geographic coordinates, elevation, date of collection (month indicated in lower case Roman numerals), habitat information, collector, depository, and preparation. Female (♀) and male (♂) symbols indicate the sex.
In accordance with
A male (holotype) and a female paratype of each newly described species was critical point dried and subsequently glued to a needle. Then we photographed them with a Keyence VHX-2000 digital microscope. Parts of each species are illustrated in the standard views of dorsal, lateral and ventral. The head is additionally depicted in frontal view (frons being vertically) and the terminalia in caudal view. Subsequent images were edited with Adobe Photoshop and Illustrator (CS6).
The specimens referred below along with the abbreviations used in the text will be deposited in the following collections:
Kuboesphasma gen. n. is placed as sistergroup to a clade (Viridiphasma + Namaquaphasma) + the remaining Austrophasmatidae based on peptide hormone sequences (Predel, unpublished data; Fig.
Kuboesphasma can be distinguished from other mantophasmatodeans, except Minutophasma, by the washed-out and indistinct butterfly-shaped spot on the frons: in the South African Austrophasmatidae sensu
Kuboesphasma compactum
None thus far.
The generic group name Kuboesphasma is a composition from the type locality, Kuboes, which is a center of the Nama community in the Richtersveld region and the ending -phasma which is commonly used to term mantophasmatodeans. The gender is neuter.
Male. SOUTH AFRICA: Kuboes, S28°26'25", E16°59'44", 18.viii.2012, 250 m, R. Predel, specimen in ethanol. Table
SOUTH AFRICA: Richtersveld S28°26'25", E16°59'44", 18.viii.2012: four males and two females, specimens in ethanol. Table
Overview of the type material including gender, locality, collection date and museum where it is deposited. Abbreviations:
Type | Gender | Locality | Collection date | Deposition | |
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Kuboesphasma compactum | Holotype | ♂ | S28°26'25", E16°59'44" | 18.viii.2012 |
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Paratype | ♂ | S28°26'25", E16°59'44" | 18.viii.2012 |
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Paratype | ♂ | S28°26'25", E16°59'44" | 18.viii.2012 |
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Paratype | ♂ | S28°26'25", E16°59'44" | 18.viii.2012 |
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Paratype | ♂ | S28°26'25", E16°59'44" | 18.viii.2012 |
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Paratype | ♀ | S28°26'25", E16°59'44" | 18.viii.2012 |
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Paratype | ♀ | S28°26'25", E16°59'44" | 18.viii.2012 |
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Minutophasma richtersveldense | Holotype | ♂ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♂ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♂ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♂ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♂ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♀ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♀ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♀ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♀ | S28°47'26.94", E17°16'20.34" | 06.ix.2014 |
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Paratype | ♂ | S28°46'31.50", E17°11'12.18" | 06.ix.2014 |
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Paratype | ♂ | S28°46'31.50", E17°11'12.18" | 06.ix.2014 |
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Paratype | ♀ | S28°46'31.50", E17°11'12.18" | 06.ix.2014 |
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Paratype | ♀ | S28°46'31.50", E17°11'12.18" | 06.ix.2014 |
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Paratype | ♀ | S28°10'20.80", E17°01'43.60" | 07.ix.2014 |
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Paratype | ♀ | S28°10'20.80", E17°01'43.60" | 07.ix.2014 |
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Measurements (male holotype followed by paratypes in parentheses, in mm): total length: 12.1 (12.5, 12.9, 12.1, 12.4); length of pronotum: 2.5 (2.6, 2.8, 2.6, 2.8); width of pronotum: 2.5 (2.4, 2.6, 2.3, 2.4); length of mesonotum: 2.0 (2.2, 2.1, 2.1, 2.3); width of mesonotum: 2.4 (2.3, 2.4, 2.2, 2.3); length of metanotum: 1.5 (1.9, 1.7, 1.7, 1.7); width of metanotum: 2.2 (2.2, 2.2, 2.1, 2.1); heights of head: 2.3 (2.3, 2.2, 2.2, 2.3); total heights of head: 2.9 (3.0, 2.9, 2.9, 3.0); width of the head: 2.8 (2.9, 3.0, 2.7, 2.9); head width over eyes: 3.0 (3.0, 3.2, 3.0, 3.0); width between eyes: 1.7 (1.7, 1.8, 1.7, 1.6); length of eye: 1.3 (1.2, 1.4, 1.4, 1.4); width of eye: 0.8 (0.8, 0.9, 0.8, 0.8).
Head (Fig.
Thorax (Fig.
Legs: green, spikes in the tibial region black; covered with setae. Prothoracic leg more massive than meso- and metathoracic ones; femur ca. three times as long as wide, with two ventro-median rows of spikes, spikes larger on pro- and mesothoracic leg, smaller on metathoracic one. Tibia green, in pro- and mesothorax ca. 10 times as long as wide, in metathoracic leg between 13 and 15 times as long, with two ventro-median rows of black spikes on pro- and mesothoracic legs, protibia with 9–12 spikes per row, on metathoracic leg only two distal spikes. Tarsus with five tarsomeres, proximal four tarsomeres with euplantulae; arolium very large.
Wings: completely absent.
Abdomen: as long as thorax and head combined; green, meso-dorsal brown longitudinal stripe weak and indistinct. Abdomen covered with setae, tergites stronger than sternits. Abdominal tergum I same width or very slightly thinner as metathorax; terga slightly broadening towards tergum VIII, terga IX and X narrowing again.
Male terminalia (Fig.
For the female only differences to the male are described. Measurements: total length: 13.5, 15.4; length of pronotum: 3.2, 3.4; width of pronotum: 3.2, 3.7; length of mesonotum: 2.0, 2.5; width of mesonotum: 3.0, 3.5; length of metanotum: 1.5, 2.0; width of metanotum: 2.8, 3.4; heights of head: 2.5, 2.8; total heights of the head: 3.2, 3.9; width of the head: 3.2, 3.8; head width over eyes: 3.4, 3.8; width between eyes: 2.1, 2.3; length of eye: 1.5, 1.7; width of eye: 0.8, 0.9.
Head (Fig.
Thorax (Fig.
Legs: protibia with 9–10 spikes per row.
Abdomen: abdomen slightly longer than head and thorax combined. No dorsal brown stripe. Widest point of abdomen at segments 5 or 6.
Female terminalia (Fig.
The species name compactum refers to the compact appearance of that species which distinguishes it easily from the second greenish species in the Richtersveld, Minutophasma richtersveldense.
Specimens were common in a heavily overgrazed area near the settlement of Kuboes; mainly in dense shrubs with small green and succulent leaves (Suaeda fruticosa, Lycium sp.).
Minutophasma gen. n. is placed as sistergroup to the Namibian genus Striatophasma.
Minutophasma can be distinguished from other mantophasmatodeans except Kuboesphasma by the washed-out and indistinct butterfly-shaped spot on the frons: in the South African Austrophasmatidae sensu
Minutophasma richtersveldense
None thus far.
The generic group name Minutophasma is a composition of the Latin word minutus that refers to the small size of that species which separates it from the other known mantophasmatodeans, and the ending -phasma which is commonly used to term mantophasmatodeans. The gender is neuter.
Male. SOUTH AFRICA: Northern Cape, north of Eksteenfontein, Richtersveld, S28°47'26.94", E17°16'20.34", 06.ix.2014, 600–700m, R. Predel, specimen in ethanol. Table
Location 1: SOUTH AFRICA, Northern Cape, north of Eksteenfontein, Richtersveld, S28°47'26.94", E17°16'20.34", 06.ix.2014, 600–700m, R. Predel: 4 males and 4 females, specimens in ethanol. Location 2: SOUTH AFRICA, Northern Cape, west of Eksteenfontein, Richtersveld, S28°46'31.50", E17°11'12.18", 06.ix.2014, 500m, R. Predel: 2 males and 2 females, specimens in ethanol. Location 3: SOUTH AFRICA, Northern Cape, Akkedis pass, Richtersveld, S28°10'20.80", E17°01'43.60", 07.ix.2014, R. Predel: 2 females, specimens in ethanol. Table
Measurements (male holotype followed by paratypes in parentheses, in mm): total length: 9.2 (location 1: 9.1, 10.5, 9.8, 9.9, 10.7) (location 2: 10.6, 10.1); length of pronotum: 1.8 (location 1: 1.9, 1.9, 1.6, 1.9, 1.6) (location 2: 2.0, 2.1); width of pronotum: 1.6 (location 1: 1.7, 1.8, 1.5, 1.8, 1,6) (location 2: 1.9, 1.8); length of mesonotum: 1.7 (location 1: 1.7, 1.7, 1.5, 1.8, 1.5) (location 2: 1.8, 1.8); width of mesonotum: 1.5 (location 1: 1.6, 1.6, 1.4, 1.7, 1.5) (location 2: 1.7, 1.6); length of metanotum: 1.3 (location 1: 1.3, 1.4, 1.4, 1.4, 1.2) (location 2: 1.4, 1.3); width of metanotum: 1.3 (location 1: 1.4, 1.5, 1.3, 1.5, 1.5) (location 2: 1.4, 1.4); heights of head: 1.7 (location 1: 1.6, 1.7, 1.5, 1.6, 1.5) (location 2: 1.7, 1.6); total heights of head: 2.1(location 1: 2.1, 2.1, 1.9, 2.0, 1.9) (location 2: 2.2, 2.2); width of the head: 2.0 (location 1: 2.0, 2.0, 1.8, 2.2, 1.8) (location 2: 2.3, 2.2); head width over eyes: 2.2 (location 1: 2.3, 2.3, 2.1, 2.4, 2.1) (location 2: 2.4, 2.4); width between eyes: 1.3 (location 1: 1.3, 1.4, 1.3, 1.3, 1.2) (location 2: 1.5, 1.5); length of eye: 1.0 (location 1: 1.0, 1.0, 1.0, 1.0, 1.1) (location 2: 1.0, 1.1); width of eye: 0.6 (location 1: 0.6, 0.6, 0.6, 0.6, 0.6) (location 2: 0.6, 0.6).
Coloration (Fig.
Head (Fig.
Thorax (Fig.
Legs: tibia with black spikes, covered with setae. Prothoracic leg more massive than meso- and metathoracic ones; profemur ca. 4 times, mesofemur ca. 4–5 times and metafemur 6–8 times as long as wide, all legs with two ventro-median rows of spikes, spikes in some specimens larger on pro- and mesothacic leg, smaller on metathoracic one. Tibia, in pro- and mesothorax ca. 8–11 times as long as wide, in metathoracic leg between 14 and 16 times as long, with two ventro-median rows of black spikes on pro- and mesothoracic legs, protibia with 5–8 spikes per row, on metathoracic leg only two distal spikes. Tarsus with five tarsomeres, proximal four tarsomeres with euplantulae; arolium very large.
Wings: completely absent.
Abdomen: as long as thorax and head combined; meso-dorsal dark longitudinal stripe strongly developed. Abdomen covered with setae. Abdominal tergum I same width or very slightly thinner as metathorax; terga slightly broadening towards tergum IX, tergum X narrowing again.
Male terminalia (Fig.
For the female only differences to the male are described. Measurements: total length: (location 1: 11.3, 12.6, 12.7, 10.7) (location 2: 11.2, 13.2) (location 3: 12.3, 12.5); length of pronotum: (location 1: 2.2, 2.4, 2.3, 2.0) (location 2: 2.2, 2.1) (location 3: 2.2, 2.3); width of pronotum: (location 1: 2.0, 2.3, 2.2, 1.7) (location 2: 2.1, 2.0) (location 3: 2.0, 2.2); length of mesonotum: (location 1: 1.9, 2.0, 2.0, 1.8) (location 2: 2.0, 1.9) (location 3: 2.0, 1.9); width of mesonotum: (location 1: 1.9, 2.1, 2.0, 1.6) (location 2: 1.9, 1.8) (location 3: 1.8, 2.1); length of metanotum: (location 1: 1.4, 1.6, 1.6, 1.3) (location 2: 1.3, 1.5) (location 3: 1.5, 1.4); width of metanotum: (location 1: 1.8, 2.0, 1.8, 1.6) (location 2: 1.8, 1.8) (location 3: 1.7, 2.0); heights of head: (location 1: 1.6, 2.1, 2.1, 1.8) (location 2: 1.9, 2.1) (location 3: 1.8, 2.1); total heights of the head: (location 1: 2.4, 2.7, 2.7, 2.4) (location 2: 2.6, 2.7) (location 3: 2.4, 2.7); width of the head: (location 1: 2.4, 2.5, 2.4, 2.2) (location 2: 2.4, 2.4) (location 3: 2.3, 2.4); head width over eyes: (location 1: 2.6, 2.7, 2.8, 2.4) (location 2: 2.7, 2.7) (location 3: 2.5, 2.5); width between eyes: (location 1: 1.6, 1.7, 1.6, 1.4) (location 2: 1.7, 1.7) (location 3: 1.7, 1.7); length of eye: (location 1: 1.1, 1.2, 1.1, 1.0) (location 2: 1.2, 1.1) (location 3: 1.1, 1.2); width of eye: (location 1: 0.6, 0.7, 0.6, 0.6) (location 2: 0.7, 0.6) (location 3: 0.7, 0.7).
Coloration: all found females are green, without dorsal longitudinal dark stripe.
Head (Fig.
Thorax (Fig.
Legs: protibia with 6–9 spikes per row.
Abdomen: no dorsal brown stripe. Widest point of abdomen at segments 5 or 6.
Female terminalia (Fig.
The species name richtersveldense refers to the currently known area of distribution, the Richtersveld.
This species was found in a variety of green and grey-green bushes with small leaves; several specimens were also collected from grass stalks. From the most northern population at Akkedis pass, only females or female nymphs (>20) were recorded over a period of three years. The absence of males implies that parthenogenesis might occur; this phenomenon was not reported from mantophasmatodeans so far. The sex ratio in the populations around Eksteenfontein was about 1:1 as usual in Mantophasmatodea.
The Richtersveld region is part of the Great Escarpment and an exceptional center of endemism within the Succulent Karoo (
The comparatively high diversity of Mantophasmatodea in the Richtersveld stands in strong contrast to other regions within their currently known distribution, where even sympatry of two species is rare (
The authors would like to thank Hans Pohl (Jena, Germany) for technical and graphical help and Siegfried Werth (Köln) for help with habitus photography (Fig.