Research Article |
Corresponding author: Ulrike Aspöck ( ulrike.aspoeck@nhm-wien.ac.at ) Academic editor: Dominique Zimmermann
© 2014 Ulrike Aspöck, Susanne Randolf.
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:
Aspöck U, Randolf S (2014) Beaded lacewings – a pictorial identification key to the genera, their biogeographics and a phylogentic analysis (Insecta: Neuroptera: Berothidae). Deutsche Entomologische Zeitschrift 61(2): 155-172. https://doi.org/10.3897/dez.61.8850
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The present paper comprises the first illustrated key for the genera of the Berothidae. Distribution maps for all genera are provided and distribution areas are discussed. A phylogenetic analysis based on the matrix of morphological characters of
Berothidae , genera, identification key, distribution maps, biogeography, cladistic analyses, phylogeny
The Berothidae Handlirsch, 1908, are a small neuropteran family that comprises about 110 known species in 24 genera. The adults superficially resemble hemerobiids (Fig.
The fossil record of Berothidae dates back as far as the Middle Jurassic, but the family is assumed to be much older (
The biology of the family is poorly documented. Since the adults are almost exclusively collected at light they are assumed to be nocturnal (
Larvae are known only from four genera of the subfamily Berothinae (Lomamyia, Isoscelipteron Costa, 1863, Podallea, Spermophorella Tillyard, 1916) and from two genera of the subfamily Nyrminae (Nyrma Navás, 1933, Berothimerobius Monserrat & Deretzky, 1999) (
The first modern treatment of the family Berothidae was completed by
The first computerized cladistic analysis of the Berothidae (
The Rhachiberothinae were primarily described as a subfamily of the Berothidae (
The present approach is based on the matrix of
Berothidae occur throughout most biogeographical regions. Cyrenoberothinae are known from southern South America and southern Africa, Nosybinae from the Afrotropical region and South America, Berothinae from most parts of the world with a high diversity in Australia and Africa (
Until now the only identification keys available are for the species of a single genus (e.g.
Figures
Localities were taken from original literature (Supplementary file
The phylogenetic analysis comprises 50 characters for 24 genera and 5 outgroup taxa. The character matrix of
Only informative characters were included in the data matrix. The cladistic analyses were performed with TNT (
A list of the currently known valid genera and species of Berothidae is provided (Supplementary file
AUSTRALIA and NEW ZEALAND (Figs
1 | New Zealand | Protobiella |
– | Australia + Barrow Island | 2 |
2 | Wings slender, proximally hardly smaller than distally (Fig. |
Stenobiella |
– | Wings proximally clearly smaller than distally (Figs |
3 |
3 | Forewings heavily falcate, gradate row in hindwings with 9–10 cross veins (Fig. |
Isoscelipteron |
– | Forewings sinuate (Fig. |
4 |
4 | Both fore and hindwings sinuate (Fig. |
Trichoma♂ |
– | Both fore and hindwings not sinuate | 5 ♀, 10 ♂ |
5 | Tergite 9 in ♀ with pseudohypocaudae (Fig. |
Austroberothella |
– | Tergite 9 in ♀ without pseudohypocaudae | 6 |
6 | Gonocoxites 9 in ♀ lacking hypocaudae (Fig. |
Trichoberotha |
– | Gonocoxites 9 in ♀ with hypocaudae (Figs |
7 |
7 | Gonocoxites 9 in ♀ with short hypocaudae (Fig. |
Trichoma |
– | Gonocoxites 9 in ♀ with long hypocaudae (Fig. |
8 |
8 | Pronotum in ♀ with brush of scales (Fig. |
disseminata group of Spermophorella |
– | Pronotum in ♀ without brush of scales | 9 |
9 | Bursa copulatrix in ♀with coiled basal element (Fig. |
Quasispermophorella |
– | Bursa copulatrix in ♀ without coiled basal element (Fig. |
goobita group of Spermophorella |
10 | Sternite 9 in ♂ with spectacular spines (Fig. |
Austroberothella |
– | Sternite 9 in ♂ without spectacular spines | 11 |
11 | Gonocoxite complex 10 in ♂ without loops (Fig. |
Trichoberotha |
– | Gonocoxite complex 10 in ♂ with loops (Figs |
12 |
12 | Gonocoxite complex 10 in ♂ with few loops (Fig. |
Spermophorella |
– | Gonocoxite complex 10 in ♂ with up to 8 screwed loops (Fig. |
Quasispermophorella |
Oriental region (Figs
1 | Forewing falcate (e.g. Fig. |
2 |
– | Forewing not falcate or sinuate | 3 |
2 | Forewing: pterostigma inconspicuous (e.g. Fig. |
Isoscelipteron |
– | Forewing: pterostigma dark, distally enlarged (Fig. |
Berotha |
3 | Postocular region globular (Fig. |
Lekrugeria |
– | Postocular region not globular (Fig. |
Nodalla |
Palaearctic region (Figs
1 | Frons elongated (Fig. |
Nyrma |
– | Frons not elongated (Fig. |
2 |
2 | Forewings falcate (e.g. Fig. |
4 |
– | Forewings not falcate | 3 |
3 | Postocular region globular (Fig. |
Lekrugeria |
– | Postocular region not globular (Fig. |
Nodalla |
4 | Length of scapus 7–10 flagellomeres (Fig. |
Podallea |
– | Length of scapus 4–5 flagellomeres (Fig. |
5 |
5 | Hindwings: gradate row with up to 10 cross veins (e.g. Fig. |
Isoscelipteron |
– | Posterior part of the hindwings without gradate row (e.g. Fig. |
Asadeteva |
Afrotropical region (Figs
1 | Forewings falcate (e.g. Fig. |
2 |
– | Forewings not falcate | 3 |
2 | Scapus length 7–10 flagellomeres (Fig. |
Podallea |
[females: spectacular spermatheca complex (Fig. |
||
– | Scapus length 4 flagellomeres (Fig. |
Berlekrumyia |
[females: spermatheca complex simple (Fig. |
||
3 | Frons elongated (Fig. |
Manselliberotha |
– | Frons not elongated (Fig. |
4 |
4 | Postocular region not globular (Fig. |
Nodalla |
– | Postocular region globular (e.g. Fig. |
5 |
5 | Costal field of forewing not enlarged (Fig. |
Tanzanberotha |
– | Costal field of forewing enlarged (Figs |
6 |
6 | Forewing with gradate row (Fig. |
Lekrugeria |
– | Forewing without gradate row (Fig. |
Nosybus |
Nearctic region, Neotropical region (Figs
1 | Forewing venation reticulate (Fig. |
Ormiscocerus/Berothimerobius |
– | Forewing venation not reticulate | 2 |
2 | Frons elongated (Fig. |
Cyrenoberotha |
– | Frons not elongated (Fig. |
3 |
3 | Forewing with 1 radial crossvein (Fig. |
Spiroberotha |
[female: hypocaudae long, finger like (Fig. |
||
– | Forewing with more than 1 radial crossvein | 4 |
4 | Forewing humeral vein recurrent, apex not falcate (Fig. |
Naizema |
[female: hypocaudae short, wart like (Fig. |
||
– | Forewing humeral vein not recurrent, apex falcate (Fig. |
Lomamyia |
[female: hypocaudae long, finger like (Fig. |
1.Frons: short (0), elongate (1), see
2.Mouthparts, position: external (0), sunken into concavity of head (1), see
3.Mouthparts, shortening: no shortening (0), shortening (1), see
4.Galea, elongation: not longer than stipes (0), longer than stipes (1), see
5.Antennal scape, length: at most twice the length of the pedicellus (0), about four times the length of the pedicellus (1), about six times the length of the pedicellus (2), see
6.Pronotum, transverse furrows: present (0), absent (1), see
7.Pronotum, number of transverse furrows: one (0), two (1), see
8.Pronotum, length: about as long as broad (0), shortened (1), elongate (2), see
9.Thorax, scales: absent (0), present (1), see
10.Forelegs: cursorial (0), raptorial (1), see
11.Wings, apex: rounded (0), subfalcate to slightly falcate (1), strongly falcate (2), see
12.Pterostigmal region: distinct (0), absent (1), see
13.Forewing, recurrent humeral vein: present (0), absent (1), see
14.Forewing, stems of R and M: separate to base (0), fused at or before base (1), see
15.Forewing, vein “b”, proximal of MP fork: present (0), absent (1), see
16.Forewing, number of sc-r crossveins: 0–2 (0), 3–5 (1), see
17.Forewing, number of CuA branches: one (0), two or more (1), see
18.Forewing, A1 vein, distal branching: with simple dichotomous marginal branching (0), additional marginal branching (1), “creeping” (=pectinate, with only short veinlets along wing margin) (2), no branching (3), see
19.Hindwing, stems of R and M: free (0), fused at base (1), see
20.Hindwing, vein “b” (free basal part of MA): sinuate (0), crossvein-like (1), absent (2), see
21.Hindwing, base of CuP: present (0), absent (1), see
22.Hindwing, distal part of CuP: simply forked (0), “creeping” (=pectinate, with only short veinlets along wing margin) (1), absent (2), see
23.Female: wings, scales: absent (0), present (1), see
24.Male: tergite 9 and ectoproct: free, not fused (0), fused (1), see
25.Male: sternite 9, caudal region: unpaired (0), paired (1).
26.Male: size of sternit 9 compared with sternite 8: of similar lenght (0), reduced (1), enlarged (2), see
27.Gonocoxit complex 11, dorsocaudal protrusion: absent (0), present (slender) (1), present (prominent) (2), see
28.Gonocoxit complex 11, fusion with gonocoxites 9: gonocoxite complex 11 associated with gonocoxites 9, but clearly separate (0), gonocoxite complex 11 partially fused with gonocoxites 9 (1), gonocoxite complex 11 amalgamated with gonocoxites 9 (2), gonocoxite complex 11 associated with ectoproct (3), see
29.Male: ninth gonocoxites, anterior apodeme: absent (0), present (1), see
30.Gonocoxit complex 10, lateral sclerites: absent (0), present (1), see
31.Gonocoxit complex 10, bristles: absent (0), present (1), see
32.Gonocoxit complex 10, length and arrangement of bristles: short and scattered (0), long and bundled (1), see
33.Gonocoxit complex 10, formations of bristles: simple bow (0), looped (1), thread-like (2), straight (3), see
34.Torulus (sclerite dorsally above the gonocoxite complex 11): absent (0), present (1), see
35.Female: tergite 9 and ectoproct: separate (0), fused, see
36.Female: tergite 9, ventral extremities: continuous with dorsal arch (0), divided from dorsal arch (1), see
37.Pseudohypocaudae: absent (0), present (1), see
38.Hypocaudae: absent (0), short (1), long (2), extremely long (3), see
39.Female: sternite 7: semiannular, posterior margin transverse (0), bilobed, posterior margin distinctly emarginate (1), consisting of a pair of lateral sclerites (gonocoxites 7) (2), see
40.Female: posterior region of sternum 7, pair of sclerite disks (gonapophyses 7): absent (0), present (1), see
41.Female: sternite 8: well developed, reaching tergite 8 laterally (0), reduced, not reaching tergite 8 laterally (1), obliterated (2), see
42:Female: sternite 8, ventral processus: absent (0), present, one pair processus (1), present, one median processus (2), see
43.Pudiculum: absent (0), present, well developed (1), present, reduced (2), see
44.Bursa copulatrix, size: small (0), large (1), see
45.Bursa copulatrix, sclerotization: membranous (0), sclerotized (1), see
46.Receptaculum seminis, globular element: globular (0), elongate (1), see
47.Ductus receptaculi: short (0), elongate (1), coiled (2), see
48.Foretarsus, male/female: 5-segmented in male and female (0), 4-segmented in male, 5-segmented in female (1), 4-segmented in male and female (2), see
49.Long ovipositor: absent (0), present (1), see
50.Forecoxae of males: not inflated (0), inflated (1).
The analysis using equal weights yielded twenty most parsimonious trees (MPTs) with a length of 209 steps, a consistency index (CI) of 0.34, and a retention index (RI) of 0.6. Bremer Support values and Bootstrap values are indicated in Fig.
Three different topologies were retrieved by implied weighting with K3-15, none of which represents a MPT. Weighting with K3-4 and K6 yielded a tree that is 212 steps long, as did weighting with K5. These two tree topologies differ only in the position of Stenobiella which is the sister group of Nodalla + Asadeteva under K5, and sister group of a larger clade comprising Nodalla + Asadeteva as well as Podallea and the Isoscelipteron clade. The two tree topologies are consistent in that Podallea is the sister group of the Isoscelipteron clade, and in Trichomatinae being the sister group to Nosybinae + Berothinae. Implied weighting with K7-15 yielded a tree with 210 steps. In that tree, the subfamily Nosybinae was not retrieved as monophyletic since the nosybinae genus Naizema was the sister group of the Trichomatinae as in all MPTs.
Schematic drawings with the identifying characteristics in the key. 2. Stenobiella cardaleae ♂, fore and hind wing; 3. Isoscelipteron rufum ♂, fore and hind wing; 4. Trichoma gracilipenne ♂, fore and hind wing; 5. Austroberothella rieki ♀, fore and hind wing; 6. Austroberothella rieki ♀, genital sclerites, lateral; 7. Trichoberotha ferruginea ♀, genital sclerites, lateral; 8. Trichoma gracilipenne ♀, genital sclerites, lateral; 9. Spermophorella goobita ♀, genital sclerites, lateral; 10. Spermophorella sp. ♀, head and pronotum lateral; 11. Quasispermophorella ingwa ♀, complex of bursa copulatrix and spermatheca; 12. Spermophorella goobita ♀, complex of bursa copulatrix and spermatheca; 13. Austroberothella rieki ♂, sternite 9 ventral; 14. Trichoberotha ferruginea ♂, gonocoxite complex 10; 15. Spermophorella kurtbaueri ♂, gonocoxite complex 10; 16. Quasispermophorella ingwa ♂, gonocoxite complex 10; 17. Berotha indica ♀, fore and hind wing; 18. Lekrugeria koenigi ♀, head dorsal; 19. Nodalla saharica, head dorsal; 20. Nyrma kervillea ♀, head frontal; 21. Nyrma kervillea ♂, forewing; 22. Podallea vasseana, ♀, head lateral; 23. Isoscelipteron fulvum, ♀, head lateral; 24. Asadeteva vartianorum ♂, hind wing; 25. Podallea arabica ♀, complex of bursa copulatrix and spermatheca; 26. Podallea squamulata ♂, gonocoxite complex 10; 27. Berlekrumyia africanella ♂, head lateral; 28. Berlekrumyia africanella ♀, complex of bursa copulatrix and spermatheca; 29. Berlekrumyia africanella ♂, gonocoxite complex 10.
Schematic drawings with the identifying characteristics in the key. 30. Manselliberotha neuropterologorum ♀, head frontal; 31. Nosybus minutus ♀, head frontal; 32. Tanzanberotha hirsuta, ♂, fore wing; 33. Lekrugeria koenigi ♀, fore wing; 34. Nosybus nobilis ♂, forewing; 35. Ormiscocerus nitidipennis ♂, forewing; 36. Cyrenoberotha penai, head frontal; 37. Spiroberotha sanctarosae ♂, head frontal; 38. Spiroberotha sanctarosae ♀, forewing; 39. Spiroberotha sanctarosae ♀, genital sclerites, lateral; 40. Spiroberotha sanctarosae ♂, gonocoxite complex 10; 41. Naizema mendozina ♂, forewing; 42. Naizema mendozina ♀, genital sclerites, lateral; 43. Naizema mendozina ♂, gonocoxite complex 10; 44. Lomamyia flavicornis, forewing; 45. Lomamyia latipennis ♀, genital sclerites, lateral; 46. Lomamyia latipennis, ♂, gonocoxite complex 10.
The distribution areas of berothid genera are extremely diverse. At present they cannot be assigned to known general patterns, although they certainly contribute to an understanding of their biogeography.
The Australian region (Figs
The Oriental region (Figs
The Palaearctic region (Figs
The Afrotropical region (Figs
Berothidae of the Nearctic and Neotropical regions (Figs
In summary, it can be stated that the distribution patterns of berothid genera are extremely heterogeneous. Their origin, biogeographic and biological backgrounds and their congruence with general patterns are far from being understood.
1) Two types of Gondwanean patterns are distinguishable: Neotropical / Afrotropical vicariance on the one hand, represented e.g. by Cyrenoberotha / Manselliberotha (Figs
2) One genus, Lekrugeria, marks a sub-Saharan Palaearctic belt which extends to the Palaearctic/Oriental transgression zone (Fig.
3) The genus with the largest and highly vicariant distribution area, Isoscelipteron, comprises the Palearctic, Oriental and Australian regions (Fig.
4) Finally: Endemism is a very common phenomenon among Berothidae, with radiations of genera within biogeographic realms, on one hand, or as monotypic enigmatic locus typicus relics, on the other hand.
The results of our cladistic analyses will be discussed in the context of previous results reported in
Generally, it is noteworthy that the trees retrieved with strong implied weighting (K3-6) are mostly congruent with the previous analysis by Aspöck and Nemenschkal (1998), while major conflicts emerge in the MPTs and the trees with implied weighting above K7. Conflicts in the shorter trees (MPTs and K7+) concern the Trichomatinae (Fig.
Within the Nosybinae the genus Nosybus (comprising seven species) and the monotypic Tanzanberotha, both from the Afrotropics (Fig.
The Berothinae represent the largest clade of Berothidae and emerged as a monophylum in all trees (Figs
One character complex that is highly interesting in the context of Berothinae and Nosybinae phylogeny is a conspicuous formation of the female sternal region of the abdominal segment seven (chars. 39 and 40): Hairy domed lateral sclerites, interpreted as gonocoxites seven, accompanied by smooth and weakly sclerotized discs which are interpreted as gonapophyses seven (
The Nyrminae (Fig.
Preferred tree of 20 most parsimonious trees produced by exhaustive search under equal weights with unambiguous character optimization (209 steps, CI = 0.34, RI = 0.6). Black circles indicate unique synapomorphies, white circles homoplastic ones. BER – Berothinae, CYR – Cyrenoberothinae, NOS – Nosybinae, NYR – Nyrminae, PRO – Protobiellinae, TRI – Trichomatinae.
The discussion of the distribution areas of the genera on a worldwide scale serves as a brain storming excerise for a forthcoming biogeographic analysis which would be premature at the present state of berothid phylogeny. Nevertheless, we expect a positive cybernetic aspect in presenting both approaches. That our phylogenetic tree should be understood as starting point for falsification does not need to be emphasized.
We greatfully acknowledge Harald Bruckner (NHMV) and Franziska Denner (NHMV) for the data acquisition and Harald Bruckner additionally for the drawings of the distribution maps, Dominique Zimmermann (NHMV) for her help with tricky TNT problems, Wolfgang Brunnbauer (NHMV) for acquisition of literature, Horst Aspöck (Medical University of Vienna) for permanent discussion on berothids and valuable comments on the manuscript, and John Plant for linguistic improvements and critically reading the manuscript. We want to thank the two reviewers for their essential input and the Museum für Naturkunde Berlin for publishing this article in open access and free of charge.
Table S1. Figure numbers in present work and in references that served as template.
Data type: references list
Sources of records upon which the distribution maps are based.
Data type: distribution data
Data table.
Data type: species data
NEXUS file.
Data type: NEXUS file
List of genera and species of Berothidae.
Data type: species list