Seven new Tettigoniidae (Orthoptera) and a new Blattellidae (Blattodea) from the Durmitor area of Montenegro with notes on previously known taxa
Author
Ingrisch, Sigfrid
Author
Pavićević, Dragan
text
Zootaxa
2010
2565
1
41
journal article
10.5281/zenodo.197230
73882c0e-4d4c-4800-bfec-0fd95f877e0e
1175-5326
197230
Poecilimon
affinis-ornatus
group
Figs. 6–12
;
Tabs. 5–6
Three forms of the
Poecilimon
affinis-ornatus
group occur in the Durmitor area. Although distributions differ, their ranges overlap and either two of the forms can occur sympatrically. It results that those three forms, despite great morphological similarity belong to three different species.
To validate similarity or differences between the taxa found on Durmitor and the nominate species of the group, cluster analyses were performed comparing the taxa with regard to (A) morphometrical data and indices, (B) characters of the stridulatory vein, (C) a combination of both, and (D) song characteristics (
Fig. 6
). However, one has to consider that cluster-analysis does not show any phylogenetic relations but simply the similarity of characters between the populations studied.
One of the forms resembles
P. o r n a t u s
(Schmidt, 1849) while two of them run to
P. a f f i n i s
(Frivaldsky, 1867) using the key in
Harz (1969)
. Indeed, in general morphology, the first form is such similar to
P. o r n a t u s
from the Alps and northern
Italy
(
Fig. 6
[as
P. pseudornatus
];
Tab. 5
) that it can only be recognised by stridulation, a syllable being more than nine times longer than in
P. o r n a t u s
(
Tab. 6
).
P. ornatus
was described from the southeastern Alps (Steiner Alps now Kamnik in
Slowenia
). Specimens at hand from Obir in Carinthia that is close to the
type
locality and from Mt. Stivo in Trentino have the stridulatory file with a distinct swelling or bump at the transient zone between the narrow apical and the large central teeth thus that the central and the apical parts forming an obtuse angle against each other (
Fig. 10
A; more distinct when using light microscopy than with this SEM micrograph!). The swelling can be regarded a characteristic feature of nominate
P. o r n a t u s
. The populations from Durmitor lack that bump on the stridulatory file (compare
Figs. 9
A and 10A). As stridulatory behaviour is of isolating value in the genus
Poecilimon
, this form obviously represents a new, sibling species. For its similarity with
P. ornatus
, we call it
Poecilimon pseudornatus
sp. n.
This new species was usually found on the high plateau of Durmitor between ca NN+
1000–1600m
; another conspecific population lives on Tara Mountain in western
Serbia
. It is possible that the new species also occurs in
Macedonia
where we found two forms similar to
P. o r n a t u s
: in one form the sinuate curvation of the stridulatory file is present, it thus obviously belongs to
P. o r n a t u s
as also
Heller (1984
,
1988
) found no distinct difference in stridulation between
P. ornatus
from the southern Alps and from
Macedonia
. In the other form the bump on the stridulatory file is absent; the fastigium verticis is however strikingly broader then in
P. pseudornatus
from
Montenegro
. Investigation of the stridulatory behaviour is necessary to settle the taxonomic status of the second form from
Macedonia
.
The two forms similar to
P. a f f i n i s
found on Durmitor differ from each other in morphology and stridulation (
Fig. 6
as
P. a. dinaricus
and
P. nonveilleri
;
Tabs. 5–6
). The first is more slender with the pronotum of the male longer than broad and the ovipositor somewhat shorter. The fastigium verticis, however, although smaller than the scapus width is comparatively broader than in the second form. The second form is stouter but has a very narrow fastigium verticis. The pronotum of the male is distinctly widening behind and varies from slightly narrower to slightly broader than long. The ovipositor is longer and more robust than in the first form. The number of teeth on the stridulatory vein is distinctly lower and the duration of a syllable shorter in the first than in the second form.
The question arises if either of both forms belongs to
P. affinis
(Frivaldsky, 1867)
. This species was described from Mehadia, in the Carpathian mountains of
Romania
. It was regarded a mountain species typical for the southern Carpathian mountains (
Kis 1962
).
Harz (1969)
lists further localities from the Pirin and Rila mountains in
Bulgaria
, from
Serbia
,
Bosna
,
Montenegro
and
Albania
.
Karaman (1974)
, comparing specimens from
Yugoslavia
with those from
Romania
, reduced the status of
P. poecilus
Ramme, 1951
(found on Šar Planina at the Serbian-Macedonian border) to a subspecies of
P. affinis
and described two new subspecies from south-western
Serbia
:
P. a. serbicus
from the environs of Priština and
P. a. hajlensis
from Hajla Planina. Another similar species,
P. komareki
Čejchan, 1957
, was described from
Albania
(environs of Tirana), and a subspecies of this,
P. k. r u m i j a e
Karaman, 1972
, from the Montenegrin coast (Mt. Rumija). In his monograph on the bioacoustics of the genus
Poecilimon
,
Heller (1984)
synonymised
P. poecilus
Ramme
with
P. a. affinis
and considered
P. k o m a re k i
as subspecies of
P. affinis
(the subspecies
P. k. rumijae
is not mentioned in his work). However, the author did not investigate specimens from
type
locality of
P. affinis
, but material from
Kosovo
,
Macedonia
and northern
Greece
. By that time, the species was not known from
Greece
according to the thorough morphological investigations of
Willemse (1982
,
1984
). But later,
Willemse (1985)
adopted the opinion and data of
Heller (1984)
. In contrast, we think it is obvious that several sibling species or subspecies similar to
P. affinis
with a rather limited distribution exist on the Balkan Peninsula.
TABLE 5.
Morphometrical data (in mm) of
Poecilimon
species of the
affinis
/
ornatus
-group [minimum-maximum (mean)]. Localities of the material studied:
P. o r n a t u s:
Mt.Stivo/Italy, Obir/Austria;
P. pseudornatus:
Durmitor, Tara Mt.
;
P. a. affinis:
Mehadia
/Romania (RO);
P. affinis
:
eastern Serbia (SR);
P. a. dinaricus:
Durmitor
;
P. nonveilleri:
Durmitor
;
P. rumijae:
Petrovaċki Strane
/Montenegro.
| Species |
Scapus |
Fastigium |
Pronotum |
Pronotum width |
Postfemur |
|
P. o r n a t u s
3
|
0.81-0.94 (0.87) |
0.97-1.09 (1.03) |
6.8-7.9 (7.3) |
6.4-7.3 (6.8) |
16.5-21.5 (19.6) |
|
P. o r n a t u s
Ƥ
|
0.90-1.00 (0.96) |
1.12-1.16 (1.21) |
7.6-8.7 (8.1) |
6.6-7.1 (6.9) |
18.5-23.0 (20.6) |
|
P. pseudornatus
3
|
0.81-0.97 (0.89) |
0.87-1.19 (1.05) |
6.5-7.9 (7.3) |
5.9-7.5 (6.6) |
19.0-23.0 (21.3) |
|
P. pseudornatus
Ƥ
|
0.90-1.03 (0.95) |
0.97-1.31 (1.16) |
7.2-8.8 (8.1) |
5.9-7.1 (6.5) |
20.0-24.0 (22.4) |
|
P. a. affinis
RO 3
|
0.90-1.03 (0.96) |
0.67-0.99 (0.82) |
6.0-8.0 (7.0) |
5.5-7.4 (6.6) |
18.0-22.0 (20.3) |
|
P. a. affinis
RO Ƥ
|
0.94-1.07 (1.01) |
0.83-0.99 (0.90) |
7.4-8.6 (7.8) |
5.9-7.5 (6.6) |
19.5-22.5 (20.9) |
|
P. affinis
SR 3
|
0.81-0.97 (0.88) |
0.62-0.78 (0.66) |
5.4-7.6 (6.4) |
5.1-7.3 (6.3) |
16.5-19.5 (18.5) |
|
P. affinis
SR Ƥ
|
0.88-1.00 (0.93) |
0.75-0.86 (0.80) |
6.2-7.8 (7.1) |
6.3-6.8 (6.6) |
19.0-20.0 (19.3) |
|
P. a. dinaricus
3
|
0.84-0.91 (0.87) |
0.66-0.78 (0.70) |
5.4-6.8 (6.2) |
4.8-6.1 (5.7) |
16.0-20.0 (18.2) |
|
P. a. dinaricus
Ƥ
|
0.87-0.94 (0.92) |
0.72-0.91 (0.81) |
6.6-7.6 (7.0) |
5.1-6.4 (5.8) |
18.0-21.0 (18.9) |
|
P. nonveilleri
3
|
0.87-0.97 (0.93) |
0.50-0.75 (0.62) |
5.9-8.0 (7.0) |
6.3-7.8 (7.1) |
17.0-21.0 (18.6) |
|
P. nonveilleri
Ƥ
|
0.90-1.00 (0.95) |
0.59-0.75 (0.69) |
6.8-8.2 (7.6) |
6.1-7.2 (6.7) |
18.0-21.5 (19.4) |
|
P. rumijae
3
|
1.03-1.09 (1.06) |
0.81-0.91 (0.86) |
9.5-9.8 (9.7) |
8.5-9.3 (8.9) |
23.5-24.0 (23.8) |
|
P. rumijae
Ƥ
|
1.25 |
0.94 |
11.8 |
9.4 |
25.0 |
| continued. |
| Species |
Body |
Scapus:Fastigium |
Pron.length:width |
Ovipositor |
n |
|
P. o r n a t u s
3
|
21.5-33.0 (28.2) |
0.76-0.97 (0.84) |
1.02-1.18 (1.07) |
17 |
|
P. o r n a t u s
Ƥ
|
24.0-32.0 (29.1) |
0.72-0.84 (0.79) |
1.11-1.23 (1.17) |
13.0-16.0 (14.9) |
10 |
|
P. pseudornatus
3
|
24.0-36.0 (29.5) |
0.73-1.00 (0.85) |
1.00-1.17 (1.10) |
34 |
|
P. pseudornatus
Ƥ
|
24.0-34.0 (28.9) |
0.73-0.97 (0.82) |
1.14-1.34 (1.24) |
16.0-19.0 (17.5) |
12 |
|
P. a. affinis
RO 3
|
22.0-28.0 (24.1) |
0.92-1.39 (1.18) |
0.95-1.23 (1.06) |
14 |
|
P. a. affinis
RO Ƥ
|
21.0-29.0 (24.4) |
1.04-1.24 (1.12) |
1.13-1.25 (1.18) |
14.5-18.0 (15.7) |
11 |
|
P. affinis
SR 3
|
22.0-27.0 (24.5) |
1.17-1.45 (1.34) |
0.83-1.17 (1.02) |
13 |
|
P. affinis
SR Ƥ
|
21.0-27.0 (23.3) |
1.14-1.21 (1.17) |
0.98-1.14 (1.06) |
11.5-14.5 (13.0) |
3 |
|
P. a. dinaricus
3
|
22.0-31.0 (25.5) |
1.12-1.32 (1.24) |
1.00-1.19 (1.09) |
11 |
|
P. a. dinaricus
Ƥ
|
19.0-32.0 (24.5) |
1.03-1.26 (1.15) |
1.14-1.29 (1.20) |
11.5-14.0 (12.8) |
8 |
|
P. nonveilleri
3
|
24.0-32.0 (27.9) |
1.26-1.88 (1.52) |
0.86-1.09 (0.98) |
15 |
|
P. nonveilleri
Ƥ
|
23.0-33.0 (27.4) |
1.25-1.58 (1.38) |
0.96-1.24 (1.14) |
15.0-17.0 (15.7) |
8 |
|
P. rumijae
3
|
32.0-37.0 (34.5) |
1.14-1.35 (1.25) |
1.03-1.14 (1.09) |
2 |
|
P. rumijae
Ƥ
|
34.0 |
1.33 |
1.26 |
21.5 |
1 |
We have not seen
P. k o m a re k i
but we had the opportunity to study specimens of the
type
series of
P. rumijae
studied by
Karaman (1972)
. A morphological comparison of
P. a f f i n i s
from the
type
locality (material in the collections of the Naturhistorisches Museum Wien and the Entomologisches Institut Zürich) with similar specimens collected in eastern
Serbia
(Avala, Stol Planina and Stara Planina) and
Montenegro
gave the following results (
Tab. 5
): With regard to external morphology, the specimens from Mehadia, eastern
Serbia
and the first form from Durmitor (
P. a. dinaricus
) belong to the same group, while the second form from Durmitor (
P. nonveilleri
) and
P. r u m i j a e
are distinctly different (
Fig. 6
A). When data of the stridulatory veins are used as reference, specimens of
P. affinis
from eastern
Serbia
are almost identical with those from Mehadia, while specimens from Durmitor differ by a lower number of teeth.
P. pseudornatus
,
P. nonveilleri
and
P. rumijae
are rather uniform with regard to this character. Finally, if data of both external morphology and the stridulatory vein are used (
Fig. 6
C),
P. affinis
from eastern
Serbia
and from Mehadia also appear to be most similar. It is thus likely that both belong to the same subspecies. However we will leave this question open until the song of specimens from the Carpathians will have been studied.
TABLE 6.
Characters of the male stridulatory file and songs of
Poecilimon
species of the
affinis
/
ornatus
-group
[minimum-maximum (mean)]. Abbreviations: TRL transition from narrow to wide stridulatory teeth (lower limit); TRU
do. (upper limit). Localities of the material studied as in
Tab. 5
.
Species Stridulatory teeth TRL TRU File with n
(number) obtuse angle continued.
|
P. ornatus
|
126-153 (140) |
70-80 (75) |
80-100 (85) |
yes |
2 |
|
P. pseudornatus
|
187-241 (219) |
110-150 (144) |
130-160 (157) |
no |
12 |
|
P. a. affinis
RO
|
172-219 (194) |
110-130 (120) |
130-150 (140) |
no |
4 |
|
P. affinis
SR
|
152-202 (188) |
100-140 (121 |
110-150 (138) |
no |
8 |
|
P. a. dinaricus
|
130-164 (144) |
60-90 (72) |
80-100 (89) |
no |
9 |
|
P. nonveilleri
|
200-250 (228) |
130-190 (151) |
140-200 (167) |
no |
11 |
|
P. rumijae
|
249-255 (252) |
150-170 (160) |
170-190 (180) |
no |
2 |
Species Syllable duration Pronounced final Main frequency t n
(ms) part (ms) (kHz) (˚C)
P. ornatus
12-20 (16) 0 15-17 (16.3)
22-26
4
P. pseudornatus
181-381 (269) 10-60 (29) 14-16 (15.3) 23-27
56
P.
a. affinis RO – – – – 0
P. affinis
SR 112-196 (168) 61-80 (69)
17 23-27
4
P. a. dinaricus
59-108 (83) 44-80 (60) 17 23-25
22
P.
nonveilleri 116-387 (262) 77-250 (154) 14-15 (14.7) 21-29
24
P.
rumijae – – – – 0 As stridulation of
P. a f f i n i s
from eastern
Serbia
(Avala) is different from the stridulation of insects from
Macedonia
and
Greece
as described in
Heller (1984
,
1988
) [compare
Fig. 11
C with
Figs. 7D
,
8
A in
Heller 1984
or Figs. 34E, 35D in
Heller 1988
], it is doubtful, if those insects from
Macedonia
and
Greece
belong to
P. affinis affinis
or to another subspecies or species. Considering this,
Poecilimon rumijae
Karaman, 1972
is here regarded as a separate species differing distinctly from
P. affinis
. The question on the status of
P. komareki
Čejchan, 1957
and its relation to other taxa of the group requires further studies of the stridulation with specimens from the
type
locality.
One of the forms from Durmitor, the population living in the Sušica canyon, comes close to typical
P. affinis
with regard to external morphology, but is rather remote to it with regard to the stridulatory vein and song (compared with specimens from eastern
Serbia
,
Fig. 6
D): the number of teeth is lower and the duration of the syllable shorter in specimens from Durmitor (
Tab. 6
). It thus represents a new subspecies
Poecilimon affinis dinaricus
s
sp. n.
The name refers to the distribution of the new taxon. This subspecies is not restricted to the Durmitor range as two males from Donje Bare (Sutjeska NP,
1500m
, Bosna-Herzegovina, leg. Willemse) agree completely with regard to external morphology and stridulatory vein with the specimens from Sušica and thus also belong to the new subspecies.
FIGURE 6.
Comparison of different species of the
Poecilimon affinis
/
ornatus
-group [cluster-analysis, distance metric is Euclidean distance, single linkage method (nearest neighbour) based on population means]. Populations studied:
P. affinis
(RO—Mehadia, Romania; SR—eastern Serbia),
P. a. dinaricus
(Durmitor)
,
P. ornatus
(Mt. Stivo, Italy; Obir, Austria; song parameters partly after Heller 1988),
P. pseudornatus
(Durmitor; Tara Mt.),
P. nonveilleri
(Durmitor)
,
P. rumijae
(Mt. Rumija, Montenegro). A, cluster using the morphometrical data "scapus: fastigium verticis x 100" (of male and female), "pronotum length: width at apex x 100 (of male), "postfemur length of male and female", "ovipositor length", Euclidean distance 0–20; B, cluster using data of the stridulatory vein "number of stridulatory teeth", "change from small to large teeth (lower and upper values)", "presence of a bump on the stridulatory vein" (presence = 100, absence = 0), Euclidean distance 0–100; C, combination of A and B, Euclidean distance 0–50; D, cluster using song characteristics "syllable duration", "duration of the pronounced final part of the syllable (if any)", "main frequency below 20 kHz", Euclidean distance 0–100.
FIGURE 7.
A–H,
Poecilimon pseudornatus
sp. n.
; I–Q,
Poecilimon nonveilleri
sp. n.
; R–V,
Poecilimon affinis dinaricus
s
sp. n.
; W–Z,
Poecilimon
affinis affinis
(of Brunner's type series from Mehadia/Romania). – A–B, I–J, R, W, Pronotum and tegmina of male in dorsal view; C–D, K–L, S, X, do. in lateral view; E, M–N, T, Y, fastigium verticis; F– G, O–P, U, Z, right cercus of male in dorsal view; H, Q, V, ovipositor.
FIGURE 8.
Poecilimon
rumijae
Karaman, 1972
(A–D, male paratype, E, female paratype, both from Petrovaċke Strane): A, Pronotum and tegmina in dorsal view; B, do. in lateral view; C, right cercus in dorsal view; D, fastigium verticis; E, ovipositor.
The third form of the group in the Durmitor area lives predominantly in the Canyons of Tara, Piva and Sušica. It resembles
P. r u m i j a e
with regard to the relations between scapus and fastigium verticis, the shape of the pronotum and the stridulatory vein. Size and the proportions of the body differ however strikingly between both forms. More material from the coastal area of
Montenegro
and the transitional region as well as investigations on stridulatory behaviour are necessary to settle the relations between both. At the present knowledge we regard the Durmitor form as a distinct, undescribed species of the
P. affinis-ornatus
group, which we call
Poecilimon nonveilleri
sp. n.
in honour of the late Prof. G. Nonveiller.