Corresponding author: Henrik Enghoff (
Trans-segmental serial colour patterns, i.e., colour patterns consisting of repeated elements, each of which covers several diplosegments / body rings, are described from several millipede taxa:
Millipedes have a general reputation of being discretely, not to say boringly coloured. There are, however, numerous exceptions, including such aptly named species as “the shocking pink dragon millipede” (
Millipedes may be uniformly coloured, or they may have more or less pronounced colour patterns. A very common pattern is one of transverse stripes, where each diplosegment is differently coloured on its front and hind parts. Instead of transverse stripes, there may be one or more contrastingly coloured spots on each diplosegment, or there may be one or more continuous longitudinal bands along the body.
Most millipedes have defensive glands secreting such repellant substances as hydrogen cyanide or benzoquinones (
Many millipede species have segmentally repeated colour patterns, e.g., contrastingly coloured paranota as in numerous species of
In a few cases, the colour pattern consists of repeated elements, each of which covers several diplosegments / body rings. These “trans-segmental serial colour patterns” are the subject of the present paper. The study was prompted by the discovery of
Specimens were examined from the following collections: MRAC: Muséum Royal d’Afrique Centrale, Tervuren, Belgium. ZMUC: Natural History Museum of Denmark (Zoological Museum), University of Copenhagen, Denmark. Further information was extracted from images kindly provided by colleagues and/or found on the internet.
Interpretation of postembryonic development in
Several species of the South African genus
I have examined five males and five females collected at Port St. Johns, Eastern Cape, South Africa, date unknown, M. Boddely leg. (MRAC). This sample is very homogeneous: All specimens possess 43 podous body rings and have no apodous body rings in front of the telson, i.e., they have the body ring formula 43+0+T sensu
Functionally, this pattern can intuitively be interpreted as aposematic (
Like at least most other members of the order
Anamorphic specimens of
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6+10+T | 31+4+T → | 35+4+T → | 39+3+T → | 42+0+T | |||
6+12+T | 21+5+T → | 26+5+T → | 31+5+T → | 36+4+T → | 40+3+T → | 43+0+T | |
28+5+T | 37+4+T → | 41+3+T → | 44+0+T | ||||
23+6+T | 45+0+T | ||||||
20+5+T |
Numbers of apodous body rings in the last seven anamorphic stadia of pachybolid millipedes. Data from the compilation by
Species | No. of podous rings in epimorphic spms | No. of apodous rings in last six anamorphic stadia | Ref. | |||||
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Dactylobolus bivirgatus (Karsch, 1881) | 35–39 | 4–5 | 4–5 | 4–5 | 4 | 3–4 | 2 | 1 |
Pelmatojulus ligulatus (Voges, 1878) | 54–55 | 6 | 6 | 6 | 6 | 5 | 3 | 1 |
Pelmatojulus insignis Saussure, 1860 | 53–56 | 5–6 | 5–6 | 5–6 | 5–6 | 4–5 | 3 | 1 |
Epibolus pulchripes (Gerstäcker, 1873) | 51–54 | 6 | 6 | 6 | 6 | 5 | 3 | 2 |
Crurifarcimen vagans Enghoff, 2011 | 56 | 6 | 6 | 6 | 6 | 5 | 3 | 3 |
The trans-segmental colour pattern in
In the east African spirostreptid
Colour patterns in 26 specimens of
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ZMUC 00101372 | 1 | F | 53+0+T | 12,4 | ? | B | B | B | B | B | B | B | b | b | T | |||||||||||||||||||||||||
ZMUC 00101374 | 3 | F | 52+0+T | 8,1 | ?13 | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | T | |||||||||||||||
ZMUC 00101372 | 2 | jF | 53+0+T | 7,2 | ?10 | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | T | ||||||||||||||||||
ZMUC 00101496 | jF | 51+2+T | 6,4 | 12! | B | B | B | B | B | B | B | B | B | B | B | b | B | B | B | B | B | B | B | a | a | T | ||||||||||||||
ZMUC 00101337 | M | 52+T+0 | 10,0 | ?16 | B | B | B | B | B | B | B | B | B | B | B | B | B | b | T | |||||||||||||||||||||
ZMUC 00101509 | M | 52+0+T | 11,0 | ? | B | B | B | B | B | B | B | B | B | B | B | B | B | B | T | |||||||||||||||||||||
ZMUC 00101495 | M | 52+0+T | 10,2 | ? | B | B | B | B | B | B | B | B | B | B | B | B | B | T | ||||||||||||||||||||||
ZMUC 00101344 | F | 52+0+T | 11,8 | ?16+ | b | b | b | b | b | T | ||||||||||||||||||||||||||||||
ZMUC 00101374 | 1 | F | 52+0+T | 10,8 | ?16 | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | T | ||||||||||||||||||
ZMUC 00101374 | 2 | jF | 52+0+T | 7,7 | ?15 | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | T | |||||||||||||||||
ZMUC 00101372 | 3 | M | 52+0+T | 9,4 | ? | b | b | b | b | b | b | b | b | b | b | b | T | |||||||||||||||||||||||
ZMUC 00101372 | 4 | jF | 52+0+T | 8,3 | ?14+ | B | B | B | B | B | B | B | B | B | B | B | B | B | T | |||||||||||||||||||||
ZMUC 00101343 | jF | 49+3+T | 5,2 | ?11 | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | a | a | a | T | ||||||||||||||
ZMUC 00101338 | 1 | jF | 51+0+T | 7,2 | ?14 | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | b | T | |||||||||||||||||
ZMUC 00101338 | 2 | j?F | 51+0+T | 9,5 | ?15 | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | T | ||||||||||||||||
ZMUC 00101373 | M | 51+0+T | 9,8 | ? | B | B | B | B | B | B | B | B | B | B | B | B | B | T | ||||||||||||||||||||||
ZMUC 00101338 | 3 | M | 51+0+T | 9,8 | ?16+ | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | b | b | T | |||||||||||||||||
ZMUC 00101339 | F | 51+0+T | 11,3 | ?17 | ? | B | B | B | B | B | B | B | B | B | B | ? | B | T | ||||||||||||||||||||||
ZMUC 00101497 | 1 | jF | 51+0+T | 8,1 | ? | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | B | T | ||||||||||||||||
ZMUC 00101497 | 2 | jF | 50+1+T | 6,1 | 12! | b | B | B | B | B | B | B | B | B | B | B | B | B | a | T | ||||||||||||||||||||
ZMUC 00101493 | jM | 45+3+T | 4,7 | 11 | B | B | b | B | B | B | B | B | B | B | B | B | B | a | a | a | T | |||||||||||||||||||
ZMUC 00101340 | 1 | j | 34+5+T | 2,5 | 5 | B | B | B | B | B | B | B | B | a | a | a | a | a | T | |||||||||||||||||||||
ZMUC 00101341 | j | 34+5+T | 2,5 | 5 | B | B | B | B | B | B | B | a | a | a | a | a | T | |||||||||||||||||||||||
ZMUC 00101342 | 1 | j | 34+5+T | 2,5 | 5 | B | B | B | B | B | B | a | a | a | a | a | T | |||||||||||||||||||||||
ZMUC 00101340 | 2 | j | 33+5+T | 2,5 | 5 | B | B | B | B | B | B | B | a | a | a | a | a | T | ||||||||||||||||||||||
ZMUC 00101342 | 2 | j | 29+5+T | 1,9 | 4 | B | B | B | B | a | a | a | a | a | T |
The relatively few anamorphic specimens (e.g.,
Body ring formulae in 26 specimens of
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ZMUC 00101372 | 1 | F | 30+5+T | 35+5+T | 40+4+T | 44+4+T | 48+?+T | ? |
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ZMUC 00101374 | 3 | F | 24+5+T | 29+5+T | 34+5+T | 39+5+T | 44+4+T | 48+3+T | 51+?+T |
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ZMUC 00101372 | 2 | jF | 25+5+T | 30+5+T | 35+4+T | 39+5+T | 44+4+T | 48+?+T |
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ZMUC 00101496 | jF | 24+6+T | 30+4+T | 34+5+T | 39+4+T | 43+4+T | 47+4+T |
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53+?0+T | |
ZMUC 00101337 | M | 24+5+T | 29+5+T | 34+4+T | 38+5+T | 43+4+T | 47+3+T | 50+?+T |
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ZMUC 00101509 | M | 24+5+T | 29+5+T | 34+4+T | 38+5+T | 43+4+T | 47+?+T | ? |
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ZMUC 00101495 | M | 24+5+T | 29+5+T | 34+5+T | 39+4+T | 43+4+T | 47+?+T | ? |
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ZMUC 00101344 | F | 28+5+T | 33+5+T | 38+5+T | 43+4+T | 47+?+T | ? |
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ZMUC 00101374 | 1 | F | 23+5+T | 28+5+T | 33+5+T | 38+5+T | 43+4+T | 47+?+T | ? |
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ZMUC 00101374 | 2 | jF | 23+6+T | 29+4+T | 33+5+T | 38+4+T | 42+5+T | 47+3+T | 50+2?+T |
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ZMUC 00101372 | 3 | M | 25+4+T | 29+6+T | 35+4+T | 39+4+T | 43+4+T | 47+?+T | ? |
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ZMUC 00101372 | 4 | jF | 30+5+T | 35+5+T | 40+4+T | 44+4+T | 48+?+T | ? |
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ZMUC 00101343 | jF | 25+5+T | 30+5+T | 35+5+T | 40+5+T | 45+4+T |
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52+0?+T | ||
ZMUC 00101339 | F | 24+5+T | 29+4+T | 33+5+T | 38+4+T | 42+4+T | 46+?+T |
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ZMUC 00101338 | 1 | jF | 23+5+T | 28+5+T | 33+5+T | 38+4+T | 42+4+T | 46+3+T | 49+2?+T |
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ZMUC 00101338 | 2 | j?F | 23+5+T | 28+5+T | 33+5+T | 38+4+T | 42+4+T | 46+4+T | 50+1?+T |
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ZMUC 00101373 | M | 24+5+T | 29+4+T | 33+5+T | 38+4+T | 42+4+T | 46+?+T |
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ZMUC 00101338 | 3 | M | 23+5+T | 28+5+T | 33+4+T | 37+5+T | 42+3+T | 45+3+T | 48+?+T |
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ZMUC 00101497 | 1 | jF | 24+6+T | 30+4+T | 34+4+T | 38+4+T | 42+4+T | 46+3+T | 49+?+T |
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ZMUC 00101497 | 2 | jF | 23+6+T | 29+4+T | 33+4+T | 37+5+T | 42+4+T | 46+?+T |
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51+0?+T |
ZMUC 00101493 | jM | 23+5+T | 28+4+T | 32+5+T | 37+4+T | 41+4+T |
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48+?+T | ||
ZMUC 00101340 | 1 | j | 24+5+T | 29+5+T |
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39+?+T | ||||
ZMUC 00101341 | j | 24+5+T | 29+5+T |
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39+?+T | |||||
ZMUC 00101342 | 1 | j | 24+5+T | 29+5+T |
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39+?+T | ||||
ZMUC 00101340 | 2 | j | 24+4+T | 28+5+T |
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38+?+T | ||||
ZMUC 00101342 | 2 | j | 23+6+T |
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34+?+T |
The bewildering multitude of inferred and observed segment formulae is summarized in
Inferred and observed body ring formulae in
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3 RO | 23+5+T (5) | 23+6+T (3) | 24+4+T (1) | 24+5+T (9) | 24+6+T (2) | 25+4+T (1) | 25+5+T (2) | 23-25+4-6 | |
4 RO | 28+4+T (1) | 28+5+T (6) | 29+4+T (4) | 29+5+T (7+ |
29+6+T (1) | 30+4+T (2) | 30+5+T (4) | 28-30+4-6 | |
5 RO | 32+5+T (1) | 33+4+T (2) | 33+5+T (7+ |
34+4+T (3) | 34+5+T (3+ |
35+4+T (2) | 35+5+T (3) | 32-35+4-5 | |
5 RO+1 | 37+4+T (1) | 37+5+T (2) | 38+4+T (6) | 38+5+T (4) | 39+4+T (3) | 39+5+T (2) | 40+4+T (2) | 40+5+T (1) | 37-40+4-5 |
5 RO+2 | 41+4+T (1) | 42+3+T (1) | 42+4+T (6) | 42+5+T (1) | 43+4+T (7) | 44+4+T (4) | 45+4+T (1) | 41-45+3-5 | |
5 RO+3 | 45+3+T (1+ |
46+3+T (2) | 46+4+T (1) | 47+3+T (2) | 47+4+T(1) | 48+3+T (1) | 49+3+T ( |
45-49+3-4 | |
5 RO+4 | 50+1+T (1+ |
51+2+T ( |
50-51+1-2 | ||||||
epimorphic spms | 51+0+T ( |
52+0+T ( |
53+0+T ( |
51-53+0 |
Numbers of apodous body rings in specimens of
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ZMUC 00101372 | 1 | 5 | 5 | 4 | 4 | yes | |||
ZMUC 00101374 | 3 | 5 | 5 | 5 | 5 | 4 | 3 | yes | |
ZMUC 00101372 | 2 | 5 | 5 | 4 | 5 | 4 | NO | ||
ZMUC 00101496 | 6 | 4 | 5 | 4 | 4 | 4 | NO | ||
ZMUC 00101337 | 5 | 5 | 4 | 5 | 4 | 3 | NO | ||
ZMUC 00101509 | 5 | 5 | 4 | 5 | 4 | NO | |||
ZMUC 00101495 | 5 | 5 | 5 | 4 | 4 | yes | |||
ZMUC 00101344 | 5 | 5 | 5 | 4 | yes | ||||
ZMUC 00101374 | 1 | 5 | 5 | 5 | 5 | 4 | yes | ||
ZMUC 00101374 | 2 | 6 | 4 | 5 | 4 | 5 | 3 | NO | |
ZMUC 00101372 | 3 | 4 | 6 | 4 | 4 | 4 | yes | ||
ZMUC 00101372 | 4 | 5 | 5 | 4 | 4 | yes | |||
ZMUC 00101343 | 5 | 5 | 5 | 5 | 4 | 3 | yes | ||
ZMUC 00101339 | 5 | 4 | 5 | 4 | 4 | NO | |||
ZMUC 00101338 | 1 | 5 | 5 | 5 | 4 | 4 | 3 | yes | |
ZMUC 00101338 | 2 | 5 | 5 | 5 | 4 | 4 | 4 | yes | |
ZMUC 00101373 | 5 | 4 | 5 | 4 | 4 | NO | |||
ZMUC 00101338 | 3 | 5 | 5 | 4 | 5 | 3 | 3 | NO | |
ZMUC 00101497 | 1 | 6 | 4 | 4 | 4 | 4 | 3 | yes | |
ZMUC 00101497 | 2 | 6 | 4 | 4 | 5 | 4 | 1 | NO | |
ZMUC 00101493 | 5 | 4 | 5 | 4 | 4 | 3 | NO |
Looking at the complete sequences, there are hardly two specimens with identical inferred sequences, although some come close:
23+5+T → 28+5+T → 33+5+T → 38+4+T → 42+4+T → 46+3–4+T: specimens ZMUC 00101338(1) and ZMUC 00101338(2), which have the epimorphic formula 51+0+T
23+5+T → 28+5+T → 33+5+T → 38+5+T → 43+4+T → 47+3/?+T. This pathway is followed by specimens ZMUC 00101344 (stadium 3RO missing) and ZMUC 00101374(1) which have the epimorphic formula 52+0+T
The inferrred course of anamorphosis in
In the course of anamorphosis of juliformian millipedes, the number of apodous rings usually reaches a maximum early in development and thereafter decreases monotonically (reaching 0 in hemianamorphic species like
Deviations from the unimodal pattern have been observed in a few other millipede species belonging to the family
Quite complicated and irregular trans-segmental colour patterns occur in several species of the order
Left:
The patterns in these
Transsegmental colour patterns in
A: | ||||||||||
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15 | 23 | 27 | 32 | 37 | 42 | 45 | 47 | ||
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14 | 22 | 32 | 40 | 46 | 50 | ||||
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6 | 15 | 24 | 34 | 45 | 52 | ||||
B: | ||||||||||
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8 | 4 | 5 | 5 | 5 | 3 | 2 | |||
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8 | 10 | 8 | 6 | 4 | |||||
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9 | 9 | 10 | 11 | 7 |
Thanks to Paul Marek, I have become aware of a striking photo by Steve Taylor, featuring a group of platydesmids from Costa Rica (
Unidentified platydesmids from Costa Rica, P. Marek & J.E. Bond leg. Adult specimens code: Left top:
There are 8 adult individuals on the photo. Each has 4–6 transsegmental pattern elements. Each element consists of a triangular light marking on each side of the midline, each triangle covering 3–5 pleurotergites and becoming narrower at its posterior end. The elements are separated by 2–3 unmarked pleurotergites. The anterior limit of each element is sharply marked. The anterior end of the specimens is difficult to identify on most specimens, so the absolute tergum numbers may be wrong by one or two. This is, however, less important – the important thing is the nature of the periodicity.
Beautiful photos of a
Trans-segmental colour patterns in unidentified platydesmid from Costa Rica (Fig. 5). A: Entries are limits between pattern elements, given as the number of the anteriormost pleurotergite in each pattern element. An attempt has been made to place pleurotergite numbers corresponding to the same stadium in the same column. B: entries are differences, expressed as no. of pleurotergites, between one limit and the one in front of it or, in other words, numbers of pleurotergites constituting one colour pattern element.
A: | ||||||||
specimen 1 | 12 | 19 | 26 | 33 | ||||
specimen 2 | 8 | 16 | 21 | 29 | ||||
specimen 3 | 14 | 21 | 29 | 37 | ||||
specimen 4 | 14 | 20 | 27 | 33 | ||||
specimen 5 | 14 | 20 | 25 | 31 | 37 | 42 | ||
specimen 6 | 7 | 15 | 20 | 26 | 33 | |||
specimen 7 | 7 | 14 | 21 | 28 | 35 | |||
specimen 8 | 7 | 15 | 22 | 28 | 35 | 40 | ||
B: | ||||||||
specimen 1 | 7 | 7 | 7 | |||||
specimen 2 | 8 | 5 | 8 | |||||
specimen 3 | 7 | 8 | ||||||
specimen 4 | 6 | 7 | 6 | |||||
specimen 5 | 6 | 5 | 6 | 6 | 5 | |||
specimen 6 | 8 | 5 | 6 | 7 | ||||
specimen 7 | 7 | 7 | 7 | 7 | ||||
specimen 8 | 8 | 7 | 6 | 7 | 5 |
Two variegated species of
Unidentified platydesmid from Malaysia. P. Pimvichai phot.
I have examined a sample of
Paratergal spots: clear lateral yellowish spots which occupy the middle of a paratergite (lateral ‘wing’). These spots occur in groups, and the posteriormost spot in each group is usually enlarged and clearly marks the limit of the pattern element. (This is in contrast to the patterns in
Less well-defined yellowish brown spots which occupy an area roughly corresponding to the basal third of the paratergites and is delimited vis-a-vis the lateral spots and vis-a-vis the paramedian tubercles by a small blackish area. These spots occur on all pleurotergites.
Pale tubercular spots: clear paramedian yellowish spots which occupy the two consecutive paramedian tubercles on a pleurotergite. These spots occur in an irregular pattern along the body, interrupted by the belowmentioned black spots.
Dark tubercular spots: paramedian black spots which occupy the two consecutive paramedian tubercles on a pleurotergite.
Paratergal and tubercular spots show asymmetrical patterns, see, e.g.,
Occurrence of lateral and paramedian colour components along the body of a
Lateral left | Paramedian tubercle left | Paramedian tubercle right | Lateral right | |
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Tergite no. | ||||
1 | ||||
2 | ||||
3 | ||||
4 | ||||
5 | ||||
6 | □ | □ | ||
7 | □ | ■ | ||
8 | ■ | □ | ||
9 | □ | ■ | ||
10 | ■ | □ | ||
11 | □ | ■ | ||
12 | ■ | □ | + | |
13 | □ | □ | ||
14 | + | □ | □ | + |
15 | ■ | □ | ||
16 | ■ | □ | ||
17 | ■ | □ | + | |
18 | ■ | □ | ||
19 | + | □ | ■ | |
20 | ■ | □ | + | |
21 | + | □ | □ | + |
22 | + | □ | □ | ++ |
23 | ■ | □ | ||
24 | + | □ | ■ | |
25 | + | □ | ■ | |
26 | + | □ | ■ | |
27 | □ | □ | + | |
28 | ++ | □ | □ | ++ |
29 | ■ | ■ | ||
30 | ■ | ■ | ||
31 | □ | ■ | ||
32 | + | □ | ■ | |
33 | ■ | □ | + | |
34 | + | □ | □ | + |
35 | + | □ | ■ | |
36 | + | □ | □ | + |
37 | ++ | □ | □ | ++ |
38 | ■ | ■ | ||
39 | ■ | □ | + | |
40 | + | □ | ■ | |
41 | + | □ | □ | + |
42 | + | □ | □ | + |
43 | + | □ | □ | + |
44 | ++ | □ | □ | ++ |
45 | + | □ | ■ | |
46 | + | □ | □ | + |
47 | ++ | □ | □ | ++ |
48 | ■ | ■ | ||
49 | ■ | □ | ||
50 | + | □ | ■ | + |
51 | + | □ | ■ | |
52 | + | □ | □ | |
53 | + | □ | □ | + |
54 | ++ | □ | ■ | |
55 | ■ | □ | ||
56 | + | □ | ■ | |
57 | + | □ | □ | + |
58 | + | □ | □ | + |
59 | ■ | ■ | ||
60 | ■ | □ | ||
61 | + | □ | □ | + |
62 | ||||
63 | ||||
64 | ||||
65 |
There is a strong tendency for the often enlarged posteriormost lateral spot in a group to occur on both sides of the same pleurotergite, in casu pleurotergites nos. (14), 22, 28, 37, 44, 47, [54], 58, (61) where numbers in round parentheses indicate ’groups’ of one spot, and the number in square brackets indicates asymmetry.
Pale tubercular spots (72 instances) are more frequent than dark ones (40 instances).
Pleurotergites with two dark tubercular spots are infrequent: 4 instances, compared with 21 instances of double-white and 31 instances of mixed colours.
There is a highly significant correlation (P= 5,33086E-13, chi2-test, 2×2 matrix,
Correlation between colour of tubercular spots and presence of lateral spots in a
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Lateral spot present | 53 | 1 |
Lateral spot absent | 19 | 39 |
(Follows almost automatically from previous): There is a tendency for the tubercular sports to be pale on both sides on the posteriormost pleurotergite of each lateral group. Thus, both tubercular spots are pale on pleurotergites 14, 22, 28, 37, 44, 47, 58 and 61, i.e. on all the pleurotergites mentioned in the previous statement except the dubious pleurotergite 54. The preponderance is highly significant (P ~0.0005, chi2-test, 2×2 matrix).
Colour pattern statistics for four specimens of
♀ |
♂ |
♀ |
♂ |
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Pattern on tergites | 6–61 | 7–59 | 6–61 | 6–50 |
White/black tubercular spots | 72/40 | 62/44 | 55/57 | 54/36 |
Double-black/double-white/mixed pair of tubercular spots | 4/21/31 | 4/21/28 | 15/13/28 | 8/17/20 |
White/black tubercular spot on hemitergite with lateral spot | 53/1 | 47/0 | 41/1 | 37/0 |
White/black tubercular spot on hemitergite without lateral spot | 19/39 | 15/44 | 14/56 | 17/36 |
P (chi2, 2×2 matrix) for values in rows 4 and 5 | 5,33086E-13 | 9,85096E-15 | 1,79343E-15 | 9,67137E-11 |
Position of enlarged posteriormost lateral spots in a group of spots in ten specimens of Pseudodesmus cf. variegatus ZMUC 00101498. (14), 22, 28, 37, 44, 47, [54], 58, (61) where symbols in round parentheses indicate ’groups’ of one spot, symbol in square brackets indicates asymmetry (spot only present on one side).
A: | |||||||||
♀ 65 tergites | (14) | 22 | 28 | 37 | 44 | 47 | [54] | 58 | (61) |
♂ 63 tergites | 19 | 26 | 34 | 42 | 49 | 55 | 59 | ||
♀ 63 tergites | (12) | 18 | 25 | 33 | 41 | 48 | 54 | 58 | (61) |
♂ 56 tergites | (14) | 20 | 28 | 36 | 43 | 50 | |||
♂ 46 tergites | 13 | 20 | 27 | 33 | 41 | ||||
♂ 47 tergites | 13 | 19 | 27 | 35 | |||||
♀ 44 tergites | 13 | 21 | 28 | 35 | |||||
j♂ 45 tergites | 13 | 19 | 25 | 33 | 39 | ||||
♀ 44 tergites | 13 | 18 | 24 | 30 | |||||
♀ 40 tergites | 13 | 19 | 25 | 34 | |||||
B: | |||||||||
♀ 65 tergites | 8 | 6 | 9 | 7 | 3 | 7 | 4 | 3 | |
♂ 63 tergites | 7 | 8 | 8 | 7 | 6 | 4 | |||
♀ 63 tergites | 6 | 7 | 8 | 8 | 7 | 6 | 4 | 3 | |
♂ 56 tergites | 6 | 8 | 8 | 7 | 7 | ||||
♂ 46 tergites | 7 | 7 | 6 | 8 | |||||
♂ 47 tergites | 6 | 8 | 8 | ||||||
♀ 44 tergites | 8 | 7 | 8 | ||||||
j♂ 45 tergites | 6 | 6 | 8 | 6 | |||||
♀ 44 tergites | 5 | 6 | 6 | ||||||
♀ 40 tergites | 6 | 6 | 9 |
Since the colour patterns of
Anamorphosis in
The anamorphosis of
4–8 pleurotergites are added between stadia I and II
3–8 pleurotergites are added between stadia II and III
4–8 pleurotergites are added between stadia II and IV
4–8 pleurotergites are added between stadia IV and V
3–8 pleurotergites are added between stadia V and VI
3–8 pleurotergites are added between stadia VI and VII
3–8 pleurotergites are added between stadia VII and VIII
The wide range of pleurotergal increments between stadia is consistent with the wide range of the numbers of pleurotergites which constitute a colour pattern element in the patterned platydesmids and andrognathids studied here. In other words, the colour pattern of an individual may well reflect the anamorphic pathway it has followed.
The holotype and only known specimen of
In several species of the Australian metopidiotrichid genus
Two species of unidentified
No species of
The colour patterns described above remain exceptions among millipedes. Whereas an aposematic functon of the pattern appears obvious in
Specimens were made available by Didier VandenSpiegel, Alexey Solodovnikov, Lars Vilhelmsen and Jan Pedersen. Guido Coza, Paul Marek and Piyatida Pimvichai gave access to photos of live animals. Nesrine Akkari and Nicolas Ioannou provided photos of preserved specimens. Richard Hoffman kindly agreed to help to make the genus name