Corresponding author: Ulrike Aspöck (
Academic editor: Dominique Zimmermann
The present paper comprises the first illustrated key for the genera of the
The
The fossil record of
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
The first modern treatment of the family
The first computerized cladistic analysis of the
The
The present approach is based on the matrix of
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
1 | New Zealand |
|
– | Australia + Barrow Island | 2 |
2 | Wings slender, proximally hardly smaller than distally (Fig. |
|
– | Wings proximally clearly smaller than distally (Figs |
3 |
3 | Forewings heavily falcate, gradate row in hindwings with 9–10 cross veins (Fig. |
|
– | Forewings sinuate (Fig. |
4 |
4 | Both fore and hindwings sinuate (Fig. |
|
– | Both fore and hindwings not sinuate | 5 ♀, 10 ♂ |
5 | Tergite 9 in ♀ with pseudohypocaudae (Fig. |
|
– | Tergite 9 in ♀ without pseudohypocaudae | 6 |
6 | Gonocoxites 9 in ♀ lacking hypocaudae (Fig. |
|
– | Gonocoxites 9 in ♀ with hypocaudae (Figs |
7 |
7 | Gonocoxites 9 in ♀ with short hypocaudae (Fig. |
|
– | Gonocoxites 9 in ♀ with long hypocaudae (Fig. |
8 |
8 | Pronotum in ♀ with brush of scales (Fig. |
|
– | Pronotum in ♀ without brush of scales | 9 |
9 | Bursa copulatrix in ♀with coiled basal element (Fig. |
|
– | Bursa copulatrix in ♀ without coiled basal element (Fig. |
|
10 | Sternite 9 in ♂ with spectacular spines (Fig. |
|
– | Sternite 9 in ♂ without spectacular spines | 11 |
11 | Gonocoxite complex 10 in ♂ without loops (Fig. |
|
– | Gonocoxite complex 10 in ♂ with loops (Figs |
12 |
12 | Gonocoxite complex 10 in ♂ with few loops (Fig. |
|
– | Gonocoxite complex 10 in ♂ with up to 8 screwed loops (Fig. |
|
1 | Forewing falcate (e.g. Fig. |
2 |
– | Forewing not falcate or sinuate | 3 |
2 | Forewing: pterostigma inconspicuous (e.g. Fig. |
|
– | Forewing: pterostigma dark, distally enlarged (Fig. |
|
3 | Postocular region globular (Fig. |
|
– | Postocular region not globular (Fig. |
|
1 | Frons elongated (Fig. |
|
– | Frons not elongated (Fig. |
2 |
2 | Forewings falcate (e.g. Fig. |
4 |
– | Forewings not falcate | 3 |
3 | Postocular region globular (Fig. |
|
– | Postocular region not globular (Fig. |
|
4 | Length of scapus 7–10 flagellomeres (Fig. |
|
– | Length of scapus 4–5 flagellomeres (Fig. |
5 |
5 | Hindwings: gradate row with up to 10 cross veins (e.g. Fig. |
|
– | Posterior part of the hindwings without gradate row (e.g. Fig. |
|
1 | Forewings falcate (e.g. Fig. |
2 |
– | Forewings not falcate | 3 |
2 | Scapus length 7–10 flagellomeres (Fig. |
|
[females: spectacular spermatheca complex (Fig. |
||
– | Scapus length 4 flagellomeres (Fig. |
|
[females: spermatheca complex simple (Fig. |
||
3 | Frons elongated (Fig. |
|
– | Frons not elongated (Fig. |
4 |
4 | Postocular region not globular (Fig. |
|
– | Postocular region globular (e.g. Fig. |
5 |
5 | Costal field of forewing not enlarged (Fig. |
|
– | Costal field of forewing enlarged (Figs |
6 |
6 | Forewing with gradate row (Fig. |
|
– | Forewing without gradate row (Fig. |
|
1 | Forewing venation reticulate (Fig. |
|
– | Forewing venation not reticulate | 2 |
2 | Frons elongated (Fig. |
|
– | Frons not elongated (Fig. |
3 |
3 | Forewing with 1 radial crossvein (Fig. |
|
[female: hypocaudae long, finger like (Fig. |
||
– | Forewing with more than 1 radial crossvein | 4 |
4 | Forewing humeral vein recurrent, apex not falcate (Fig. |
|
[female: hypocaudae short, wart like (Fig. |
||
– | Forewing humeral vein not recurrent, apex falcate (Fig. |
|
[female: hypocaudae long, finger like (Fig. |
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
Schematic drawings with the identifying characteristics in the key.
Schematic drawings with the identifying characteristics in the key.
Distribution maps of the genera of
Distribution maps of the genera of
Distribution map of the genera of
Distribution map of
Distribution map of the genera of
Distribution map of the genus
Distribution map of the subfamilies of
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
The
The
The
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
2) One genus,
3) The genus with the largest and highly vicariant distribution area,
4) Finally: Endemism is a very common phenomenon among
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
Within the
The
One character complex that is highly interesting in the context of
The
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 –
Strict consensus tree produced by exhaustive search under equal weights with bootstrap values over 50% (above) and Bremer support values (below).
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.
references list
Sources of records upon which the distribution maps are based.
distribution data
Data table.
species data
NEXUS file.
NEXUS file
List of genera and species of Berothidae.
species list