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Zootaxa 4808 (1): 131–140 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4808.1.7 http://zoobank.org/urn:lsid:zoobank.org:pub:82D8C720-6938-4AF2-9186-491F07053EC8

Rhachiella malawica gen. nov., spec. nov. from Malawi—another beauty of the Afrotropics (Neuroptera: Rhachiberothidae)

ULRIKE ASPÖCK1,2, HORST ASPÖCK3, JAMES B. JOHNSON4, TRUST KASAMBALA DONGA5 & PETER DUELLI6,* 1Naturhistorisches Museum Wien, Zweite Zoologische Abteilung, Burgring 7, A-1010 Vienna, Austria. �[email protected]; https://orcid.org/0000-0003-1912-2609 2Department of Evolutionary Biology, University of Vienna, Althanstraße 14, A-1090 Vienna, Austria. 3Institute of Specific Prophylaxis and Tropical Medicine, Medical University of Vienna, Kinderspitalgasse 15, A-1090 Vienna, Austria. �[email protected]; https://orcid.org/0000-0001-9407-3566 4Department of Entomology, Plant Pathology, and Nematology, University of Idaho, Moscow, ID, 83844, USA �[email protected]; https://orcid.org/0000-0002-0539-0998 5Lilongwe University of Agriculture and Natural Resources (LUANAR), Bunda College Campus, P.O. BOX 219, Lilongwe, Malawi �[email protected]; https://orcid.org/0000-0002-2445-3405 6Biodiversity and Conservation Biology, Swiss Federal Research Institute WSL, Zürcherstrasse 111, CH-8903 Birmensdorf ZH, Switzerland. �[email protected]; https://orcid.org/0000-0001-8862-8262 *Corresponding author

Abstract

A new species and a new genus of Rhachiberothidae, Rhachiella malawica gen. nov., spec. nov., are described from Malawi. The new species is characterized by a flat vertex, a long penisfilum in the male, and by a bifurcate pseudohypocauda in the female. This combination of characters requires the description of a new genus, which is the sister taxon of Mucroberotha Tjeder, 1959. This is the first record of Rhachiberothidae in Malawi. The distributions of all 14 species of Rhachiberothidae so far known are shown in three maps.

Key words: Neuropterida, Thorny Lacewings, taxonomy, distribution

Introduction

The family Rhachiberothidae Tjeder, 1959, currently comprises three genera: the closely related Rhachiberotha Tjeder, 1959, and Mucroberotha Tjeder, 1959, and their sister-taxon Hoelzeliella U. Aspöck & H. Aspöck, 1997. According to our present knowledge the family is restricted to the Afrotropics, reaching its north-eastern boundaries in Ethiopia (Figs 12–14) (Tjeder 1959, 1968; U. Aspöck & Mansell 1994, U. Aspöck & H. Aspöck 1997). Rhachiberotha with hitherto five described species is known from southern parts of the Afrotropics: R. signifera Tjeder, 1959 (Zimbabwe), R. smithersi Tjeder, 1959 (Zimbabwe), R. ingwe U. Aspöck & Mansell, 1994 (South Af- rica), R. sheilae U. Aspöck & Mansell, 1994 (South Africa), and R. pulchra U. Aspöck & H. Aspöck, 1997 (South Africa, Namibia). Mucroberotha with hitherto seven described species is known from southern and eastern parts of the Afro- tropics: M. fasciata Tjeder, 1959 (Zimbabwe), M. nigrescens Tjeder, 1968 (Zimbabwe), M. vesicaria Tjeder, 1968 (Zimbabwe, South Africa, Namibia), M. aethiopica U. Aspöck & Mansell, 1994 (Ethiopia), M. angolana U. Aspöck & Mansell, 1994 (Angola), M. minteri U. Aspöck & Mansell, 1994 (Zimbabwe, Namibia), and M. copelandi U. Aspöck & H. Aspöck, 1997 (Kenya, Tanzania). The monotypic genus Hoelzeliella with H. manselli U. Aspöck & H. Aspöck, 1997, is known from the type locality in South Africa only. The phylogenetic position of the family Rhachiberothidae—originally described as a subfamily of the Berothi- dae by Tjeder (1959), subsequently interpreted as a subfamily of the Mantispidae by Willmann (1990), then elevated

Accepted by A. Letardi: 15 May 2020; published: 1 Jul. 2020 131 to family rank by U. Aspöck & Mansell (1994) as the sister-group of Berothidae, and also treated in this position in U. Aspöck et al. (2001), U. Aspöck & H. Aspöck (2008), Beutel et al. (2010), Zimmermann et al. (2011), Randolf et al. (2014), finally interpreted as the sister-group of Mantispidae in Wang et al. (2016), and as sister-group of a part (Symphrasinae) of paraphyletic Mantispidae in Winterton et al. (2018). Oswald (2019) in his data base still keeps the subfamily status of Rhachiberothinae within the family Berothidae. The key to the genera in U. Aspöck & Mansell (1994) differentiates Rhachiberotha with „vertex of head having three tubercles; forewing with stem of R and of M free to wing base“, from Mucroberotha with „vertex of head forming one dome; forewing with stems of R and M fused at wing base“. The differentiation was upheld also in U. Aspöck & H. Aspöck (1997). However, in connection with the discovery of the new rhachiberothid species from Malawi described in the present paper this differentiation appears oversimplified, as the stems of R and M of the forewings are not fused at the wing base as in Mucroberotha, but are free as in Rhachiberotha, whereas the vertex of the head lacks three tubercles as in Rhachiberotha, and is not as typically domed as in Mucroberotha. The male genitalia are equipped with a long whip-like penisfilum, as is typical for Mucroberotha, however, the female gonocoxites 9 and the pseudohypocaudae (processes of tergite 9) look different. Taken all these arguments together the description of a new genus seems necessary.

Material and methods

The specimens examined in the present study are deposited in the W. F. Barr Entomological Museum at the Univer- sity of Idaho, Moscow, USA (WFBEMI), the Natural History Museum Vienna, Austria (NHMW), in the collection of Horst and Ulrike Aspöck, Vienna, Austria (HUAC), and in the Museums of Malawi (MM), Chichiri Blantyre 3, Malawi. Genitalic preparations were made by clearing the apex of the abdomen in KOH for five hours. After rinsing the KOH with distilled water, the apex of the abdomen was transferred to glycerine for further examination. Habitus photos were taken with a Leica DFC camera attached to a Leica MZ16 binocular microscope and processed with the help of Leica Application Suite. They were then stacked with Zerene Stacker 64-bit and processed with Adobe Photoshop Elements 8. For the distribution maps the Natural Earth program was used. Terminology of the genitalia follows that of U. Aspöck & H. Aspöck (2008).

Taxonomy

Genus Rhachiella gen. nov.

Type species (by present designation): Rhachiella malawica spec. nov.

Diagnosis. Head with flat vertex, slightly inflated genae, scape twice as long as broad. Pronotum approximately rectangular. Apices of wings rounded. Radius and Media free to wing base. Male with penisfilum; female with pseudohypocaudae having two apical processes. Description: With the characters of the type species. Systematics: Rhachiella gen. nov. is the sister-group of Mucroberotha with the long penisfilum of the males as a synapomorphy. Rhachiberotha + (Mucroberotha + Rhachiella gen. nov.) constitute the sister-group of Hoel- zeliella.

Rhachiella malawica spec. nov.

Holotype: male, Malawi, Southern Province, Lake Zomba, 1458m, -015°21’37” / +035°18’32”, 17-XI- 2018 JB Johnson, P. Duelli, T. Donga leg. Type locality: Malawi, Lake Zomba.

132 · Zootaxa 4808 (1) © 2020 Magnolia Press ASPÖCK ET AL. General appearance (Figs 1, 2, 3) with the typical rhachiberothid look, hairiness, bright yellowish habitus, with only basal parts of antennae dark brown; pterostigma of male brownish, with a yellowish patch in the centre, however yellowish with a brownish patch in the centre in the female. Length of forewing of males 6.5–6.7 mm, of females 6.8–7.2 mm.

FIGURE 1. Rhachiella malawica spec. nov., paratype, female, habitus (photo Thomas Reich, WSL Birmensdorf, Switzerland). Length of forewing 7 mm.

Description of male (Figs 2–4): Head flat, yellow, with paired brown stripes on the vertex, laterally brownish; frons yellow with brown spot, base of clypeus dark brown, genae slightly inflated; scape twice as long as broad, scape, pedicellus and first segment of flagellum dark brown, rest of flagellum yellow. Mouthparts dark brown. Pro- notum brownish, with two pairs of brownish tubercles. Meso- and metanotum with a yellow median fascia, and with a median brown line. Meso- and metanotum brown, mesonotum with yellowish patches laterally. Femora of raptorial forelegs with two rows of brown, comb-like thorns (Fig. 3). Forewing (Figs 2, 4): Radius and Media free to wing base. Pterostigma brownish, with a yellow patch. Wing membrane hyaline with light smoky shadows around cross veins and forkings. Venation: longitudinal veins yellowish, cross-veins brownish. Hindwing: Basal Media anterior sinuate. Pterostigma brownish, much smaller than in forewing. Abdomen: Tergites yellow, with median brown line, pleurae yellow with brownish zig-zag line, sternites whit- ish-yellowish. Genital segments of male (Figs 5–6): Tergite and sternite of segment 8 much shorter than the previous ones, with distinct spiraculum, tergite 9 ribbon-like with a small ridge. Sternite 9 inconspicuous, short, with deep incision. Ectoproct with distinct trichobothria rosette, apex broad finger-shaped, sculptured. Gonocoxites 9+11 extremely reduced to a tiny weakly sclerotized structure. Gonocoxites 10 paired basally, fused with the unpaired gonapophyses 10, terminating in a long penisfilum. Hypandrium internum of typical shape. Description of female (Figs 1, 7): Eidonomic characters similar to those of the male. Pterostigma yellowish with brownish patch in the centre, however less contrasting than in the male.

Rhachiella malawica gen. nov., spec. nov. from Malawi Zootaxa 4808 (1) © 2020 Magnolia Press · 133 FIGURES 2–6. Rhachiella malawica spec. nov. (photos Harald Bruckner, NHMW). 2. Paratype, male, lateral; 3. Paratype, male, head and foreleg, lateral; 4. Paratype, male, distal half of wings; 5. Holotype, male, genital sclerites, lateral; 6. Holotype, male, genital sclerites, ventral. dp: digital process of ectoproct; e: ectoproct; gx 9 + gx 11: fused complexes of gonocoxites 9 and gonocoxites 11; gx 10 + gp 10: fused complexes of gonocoxites 10 and gonapophyses 10; hi: hypandrium internum; pf: penisfilum; S: sternite; T: tergite; ta: tarsus; ti: tibia; tr: trochanter.

134 · Zootaxa 4808 (1) © 2020 Magnolia Press ASPÖCK ET AL. Genital segments of female (Figs 8–10): Sternite 7 reduced to a small band-like sclerite. Pseudohypocaudae with two apical digitiform processes. Gonocoxites 9 with short apex (hypocaudae). Bursa copulatrix connected by a long ductus with receptaculum seminis, which comprises a complex system of ducts. Rhachiella malawica spec. nov. is differentiated from all known species of Rhachiberotha by the lack of tubercles on the head, by possession of a penisfilum in the male (lacking in Rhachiberotha), and by pseudohypocaudae with two processes (only a simple apex in Rhachiberotha). Rhachiella malawica spec. nov. is differentiated from all known species of Mucroberotha by a flat head (domed in Mucroberotha), by Radius and Media of forewing free to wing base (fused in Mucroberotha).

FIGURES 7–10. Rhachiella malawica spec. nov., paratype, female (photos Harald Bruckner, NHMW). 7. Right pair of wings spread; 8. Genital sclerites, lateral; 9. Genital sclerites, ventrolateral; 10. Genital sclerites, lateral. e: ectoproct; C: Costa; gx 9: gonocoxites 9; M: Media; MA: basal piece of Media anterior; ph: pseudohypocaudae with digitiform processes (black arrows); Pt: pterostigma; rs: receptaculum seminis; Sc: Subcosta; T: tergite.

Rhachiella malawica gen. nov., spec. nov. from Malawi Zootaxa 4808 (1) © 2020 Magnolia Press · 135 Rhachiella malawica spec. nov. is differentiated from Hoelzeliella by the rounded apex of the forewing (slightly sinuate in Hoelzeliella). Etymology: Rhachiella is derived from Rhachiberotha. It has feminine gender; malawica is an adjective derived from the country where the species has been discovered. Material studied: Holotype, male, and 4 males, and 4 females, paratypes: Malawi, Southern Region, Lake Zomba, 1458m, -015°21’37” / +035°18’32”, 17 Nov 2018, J.B. Johnson, P. Duelli, T. Donga leg. Holotype, and 1 female in MM, 1 male and 1 female in WFBEMI, 2 males and 1 female in NHMW, 1 male and 1 female in HUAC. The specimens were taken 30 m from the lake shore of the Zomba-dam, mainly on trees at the edge of natural vegetation, while most other trees along the shore were ornamentals. It was noteworthy that the times of the col- lections occurred between mid-afternoon and early evening, mostly 3-5 PM local time. Trees that had not yielded specimens of R. malawica earlier in the day did yield specimens when revisited during this time period. 1 male paratype (MM): Malawi, Southern Region, Zomba, Forest path near Sunbird Hotel, 1521m, -015°21’55” / +035°18’5”, 16 Nov 2018, JB Johnson, P. Duelli, T. Donga leg., collected on a bush along a path on a steep slope. Rain forest, natural vegetation (Fig. 11). Rhachiella malawica spec. nov. is known from the region of Zomba in Malawi only and thus in the centre of the hitherto known distribution of Rhachiberothidae (Figs 12–14).

FIGURE 11. Forest habitat of Rhachiella malawica spec. nov., near Sunbird Hotel, Zomba (photo P. Duelli).

Mucroberotha copelandi U. Aspöck & H. Aspöck, 1997

New record: 1 male: “Kenia, 21.4.01 Mesusop Elas Kopje, P. Duelli leg.”, 1 female: “KENYA, Eastern Prov. Base of Ukasi Hill, 613 m, 0.82103°S, 38,54443°E / Malaise trap, Acacia / Commiphora savannah, 7-21 NOV 2011, R. Copeland”.

136 · Zootaxa 4808 (1) © 2020 Magnolia Press ASPÖCK ET AL. FIGURE 12. Distribution of Rhachiberothidae (based on U. Aspöck & H. Aspöck 1997). 1 = Rhachiberotha signifera, 2 = Rh. ingwe, 3 = Rh. smithersi, 4 = Rh. sheilae, 5 = Rh. pulchra, 6 = Mucroberotha fasciata. Map from Natural Earth.

Discussion

With the description of Rhachiella malawica altogether 14 extant species of the family Rhachiberothidae in four genera are known. Their distribution is confined to sub-Saharan parts of Africa. With the exception of Mucroberotha vesicaria these species have been recorded from very few, even from single localities, and have been taken in few specimens only. Mucroberotha vesicaria is distributed over a larger part of southern Africa (Fig. 13). This species is sometimes frequent at light traps. Also, the other species have mostly been taken at light, Rhachiella malawica, however, has been collected by sweeping vegetation in the day time. The larvae and larval biology of Rhachiberothidae are almost unknown. Eggs and first-instar larva of Mu- croberotha vesicaria Tjeder have been described by Minter (1990). First-instar larvae were fed on spider eggs, lepidopteran larvae and worker-caste termites. However, Minter did not succeed in inducing development of later instars. Mantispidae and Berothidae have—though different in each family—an unusual hypermetamorphosis during larval life. In the context of the clade constituted by Mantispidae, Berothidae and Rhachiberothidae—irrespective of their sister-group relationships—a hypermetamorphosis of the larvae may be also hypothesized for Rhachiberothi- dae. It is, however, in reality not known whether the second and third instar are parasitic, and if so, it remains still open with which hosts they are associated. The Golden Age of Rhachiberothidae was long ago. So far 17 fossil species have been described from various parts of the Old World (Baltic amber, England, France, Lebanon, Myanmar, Japan) and New World (Canada, USA: New Jersey). All fossil specimens were found in amber from various periods: Lower Cretaceous (Lebanon, France), early Upper Cretaceous (Myanmar, France), Late Cretaceous (New Jersey, Japan, Canada), Early Eocene (France), Mid-Eocene (Baltic amber, England). Two of these species are assigned to the subfamily Rhachiberothinae, 14 to the extinct subfamily Paraberothinae, one species could not be assigned to any of these two subfamilies (Engel

Rhachiella malawica gen. nov., spec. nov. from Malawi Zootaxa 4808 (1) © 2020 Magnolia Press · 137 2004; Engel & Grimaldi 2008; Grimaldi 2000; Makarkin & Kupryjanowicz 2010; Makarkin 2015; McKellar & Engel 2009; Nel et al. 2005 a, b; Petrulevičius et al. 2010; Schlüter 1978; Shi et al. 2015; Whalley 1980; 1983). Nakamine & Yamamoto (2018) give a detailed list of all fossil species.

FIGURE 13. Distribution of Rhachiberothidae (based on U. Aspöck & H. Aspöck 1997). 7 = Mucroberotha vesicaria. Map from Natural Earth.

8 = M. nigrescens 9 = M. aethiopica 10 = M. angolana 11 = M. minteri 12 = M. copelandi 13 = Rh. malawica 14 = H. manselli

FIGURE 14. Distribution of Rhachiberothidae (based on U. Aspöck & H. Aspöck 1997). 8 = Mucroberotha nigrescens, 9 = M. aethiopica, 10 = M. angolana, 11 = M. minteri, 12 = M. copelandi,13 = Rhachiella malawica spec. nov., 14 = Hoelzeliella manselli. Map from Natural Earth.

We are thankful to the National Commission for Science and Technology (Lilongwe, Malawi) for the collection permit and the Environmental Affairs Department for the export permit (Permit EAD-12-07-031-17-21). Cordial thanks to Mag. Harald Bruckner, Natural History Museum Vienna, for providing the photographs in the lab, and to Thomas Reich for the live picture of Rhachiella malawica spec. nov. The distribution maps were kindly provided by Mag. Harald Bruckner, Josef Muhsil-Schamall, and Dr. Rudolf Pavuza (NHMW)—many thanks!

References

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