Revision of the Eurybrachidae XIV. The Australian genera Olonia Stål, 1862 and Stalobrachys gen. nov. (Hemiptera: Fulgoromorpha)
Author
Constant, Jérôme
text
European Journal of Taxonomy
2018
2018-12-13
486
1
97
journal article
28761
10.5852/ejt.2018.486
0e26b0d8-14bc-46b2-aac6-feed5d62b2d8
2270151
urn:lsid:zoobank.org:pub:510A70C2-01F5-4C6E-855D-EFE140B45664
Olonia marginata
Distant, 1906
Figs 3
,
15–25
Olonia marginata
Distant, 1906: 206
(described).
Olonia marginata
–
Kirkaldy 1907: 105
(listed from
Queensland
). —
Metcalf 1956: 64
(catalogued).
Diagnosis
This species can be recognized by the following combination of characters:
(1) hind wings without orange marking (
Figs 16A
,
20A
)
(2) pro- and mesofemora and -tibiae largely black-brown (
Figs 16A–D
,
20A–D
)
(3) anal tube of male spatulate, constricted at basal ½ (
Fig. 19B
)
(4) centroventral part of gonostyli with narrow and strongly elongated process that is sinuate basally and hooked apically (
Fig. 19A, C
)
(5) laterodorsal part of gonostyli with narrow and strongly elongated process that is strongly curved posteroventrally (
Fig. 19A, C–D
)
(6) rather small size:
6–8 mm
Etymology
The species epithet
marginatus
(adjective, Latin) means ʻmarginatedʼ, ʻborderedʼ. It refers to the darkened costal and apical margins of the tegmina.
Material examined
Lectotype
AUSTRALIA
•
1 ♀
;
Queensland
; “Type”, “Type”, “
Olonia marginata
, type, Dist.”, “Queensland,
F.P. Dodd
,
1902
– 319”;
here designated
to provide a reference standard for this species;
BMNH
(
Fig. 15
).
Paralectotypes
AUSTRALIA
•
2 ♀♀
; “
Queensland
,
F.P. Dodd
,
1902
–
319
”;
BMNH
.
Note
In the collections of
BMNH
, there are 3 additional specimens identified by Distant but which are not part of the
type
series:
1 ♀
, “
Olonia marginata
Dist.
”, “Townsville [
19°25′26″ S
,
146°56′44″ E
] (Dodd)”, “B”, “Distant Coll. 1911-383”; 2 ♂♂, “
Queensland
,
F.P. Dodd
, Brit. Mus.,
1907—54
”.
These specimens are particularly interesting because (1) they are from the same collector as the
type
series; (2) they provide Townsville as the location for the
type
series, where the collector, Frederick Parkhurst Dodd (
1861–1937
), lived at the time before moving to Kuranda and (3) there are two males, which allows a characterization of the species based on male genitalia. These males are used here as a reference for the recognition of this species.
Additional material
AUSTRALIA
• 3 ♂♂,
1 ♀
;
Townsville
;
Jan. 1945
;
B. Malkin
leg.;
USNM
• 1 ♂,
1 ♀
; same collection data as for preceding;
RBINS
•
1 ♀
;
Townsville
;
2 Mar. 1956
;
I. Sutherland
leg.; QM
• 1 ♂; Townsville, Hervey Range;
19°22′42″ S
,
146°31′29″ E
;
22 Sep. 1981
; D.C. Geijskes leg.; RMNH
• 1 ♂; Townsville, Heatley; 19°17′35″ S, 146°45′09″ E;
4 Aug. 1976
;
D.C. and R. Geijskes
leg.;
RMNH
• 1 ♂;
Townsville
;
7 Apr. 1977
;
Brown
leg.;
QPIM
• 1 ♂;
Pallarenda Point
near
Townsville
;
19°11′20″ S
,
146°46′25″ E
,
26 Jan. 1965
; E.C. Dahms leg.; open forest,
by net
; QM
• 1 ♂; Townsville, Hyde Park Shopping Centre; 19°16′46″ S, 146°47′49″ E;
25 Feb. 1992
;
L.M. Brown
leg.;
on
Melaleuca quinquenervia
;
ASCU
• 1 ♂;
Macrossan
;
20°00′18″ S
,
146°26′44″ E
;
15 Apr. 1958
;
K.L. Hartley
leg.;
ANIC
• 1 ♂; near
Ingham
;
18°39′00″ S
,
146°10′00″ E
;
13 Mar. 1961
;
R. Straatman
leg.; on mangrove forest beach;
ANIC
• 2 ♂♂,
1 ♀
;
10 mi
. SSE of
Collinsville
[coordinates of
Collinsville
:
20°33′08″ S
,
147°50′38″ E
];
14 Sep. 1950
;
E.F. Riek
leg.;
ANIC
• 5 ♂♂,
1 ♀
;
30 mi
. S of
Ayr
; [coordinates of
Ayr
:
19°34′33″ S
,
147°24′18″ E
];
9 Sep. 1950
; E.F. Riek leg.; ANIC
• 2 ♂♂,
1 ♀
; same collection data as for preceding;
RBINS
• 1 ♂;
35 mi
. SE of
Ayr
;
3 Oct. 1950
;
E.F. Riek
leg.;
ANIC
•
1 ♀
;
40 mi
. SW of
Ayr
;
7 Oct. 1950
;
E.F. Riek
leg.;
ANIC
•
1 ♀
;
40 mi
. S of
Ayr
;
10 Sep. 1950
;
E.F. Riek
leg.;
ANIC
•
1 ♀
;
50 mi
. S of
Ayr
;
11 Sep. 1950
;
E.F. Riek
leg.;
ANIC
• 1 ♂;
35 mi
. SW of
Ayr
;
6 Oct. 1950
; E.F. Riek leg.; ANIC
• 1 ♂,
1 ♀
;
60 mi
. SW of
Ayr
;
8 Oct. 1950
;
E.F. Riek
leg.;
ANIC
• 1 ♂,
1 ♀
; same collection data as for preceding;
RBINS
•
1 ♀
;
35 mi
. NW of
Bowen
; [coordinates of
Bowen
:
20°00′33″ S
,
148°13′46″ E
];
2 Oct. 1950
;
E.F. Riek
leg.;
ANIC
• 2 ♂♂;
25 mi
. NW of
Bowen
;
1 Oct. 1950
;
E.F. Riek
leg.;
ANIC
• 1 ♂;
10 mi
. S of
Bowen
;
26 Sep. 1950
;
E.F. Riek
leg.;
ANIC
•
4 ♀♀
;
Townsville
; alt. sea level;
13 Jan. 1962
; E.S. Ross and D.Q. Cavagnaro leg.; CAS
• 1 ♀; same collection data as for preceding; RBINS
• 1 ♂,
1 ♀
;
Majors Creek
;
19°37′36″ S
,
146°57′28″ E
;
27 Oct. 2006
;
G. Cocks
leg.; on grapes;
RBINS
• 2 ♂♂,
2 ♀♀
;
Townsville
;
12 Oct. 1983
;
M.E. Irwin
and
E.I. Schlinger
leg.;
INHS
• 1 ♂,
1 ♀
; same collection data as for preceding;
RBINS
•
1 ♀
;
70 km
NW of
Townsville
;
9 Nov. 1990
;
W.F. Chamberlain
leg.;
TAMU
• 1 ♂,
1 ♀
;
Rollingstone
;
19°02′40″ S
,
146°23′01″ E
;
19 Apr. 1998
;
L.J. Cookson
leg.;
MVMA
• 1 ♂;
Surveyor Creek
;
20°43′59″ S
,
148°36′00″ E
;
30 Jan. 1991
;
L.J. Cookson
leg.;
MVMA
• 1 ♂,
1 ♀
;
Bowen
,
Queens Bay
;
19°58′43″ S
,
148°14′50″ E
;
7 Jul. 1971
;
Z. Liepa
leg.;
ANIC
• 1 ♂;
30 km
S of
Ingham
;
10 Nov. 1990
;
G.M. Chamberlain
leg.;
TAMU
• 1 ♂;
Clare
;
19°49′43″ S
,
147°11′46″ E
;
7 Apr. 1951
; W.
A.S.
leg.; from sweeping in jute;
QDPI
• 1 ♂;
Saltwater Creek
near
Toomulla
;
19°05′56.37″ S
,
146°27′46.69″ E
;
20 Apr. 1998
;
L.J. Cookson
leg.;
MVMA
• 1 ♂;
32 km
S of
Ayr
;
6 Feb. 1964
;
J. Sedlacek
leg.;
BPBM
• 1 ♂;
Gumlu
,
48 km
SE of Ayr
;
19°54′09″ S
,
147°35′30″ E
;
6 Feb. 1964
;
J. Sedlacek
leg.;
BPBM
• 1 ♂;
Herbert River
;
18°31′59″ S
,
146°18′00″ E
; 1919;
J.F. Illingworth
leg.; on (sugar)cane;
BPBM
• 1 ♂,
2 ♀♀
;
Townsville
,
Strand Beach
,
Northern End
; [coordinates of
Strand Beach
:
19°14′23″ S
,
146°48′24″ E
];
15 Mar. 2016
; V. Ryland leg.;
on
Ipomoea
pes-caprae; RBINS
• 2 ♀♀; same collection data as preceding;
19 Feb. 2016
; QM
• 1 ♀; same collection data as preceding;
25 Feb. 2016
; QM
• 1 ♂; Townsville, port wall; [coordinates of Townsville port: 19°15′17″ S, 146°50′11″ E];
15 Mar. 2016
;
V. Ryland
leg.;
on
Ipomoea
pes-caprae;
RBINS
• 1 ♂;
Townsville
, small boat harbour wall; [coordinates of
Townsville
small boat harbour:
19°15′12″ S
,
146°49′24″ E
];
19 Mar. 2016
; V. Ryland leg.;
on
Ipomoea
pes-caprae; RBINS
• 1 ♂; same collection data as for preceding; QM
• 1 ♂; Townsville, Bushland Beach; 19°11′19″ S, 146°40′40″ E;
4 Apr. 2016
;
V. Ryland
leg.;
on
Ipomoea
pes-caprae;
RBINS
• 1 ♂,
1 ♀
; same collection data as for preceding; QM
• 2 ♂♂;
Magnetic Island
,
Nelly Bay
Beach; [coordinates of
Nelly Bay
:
19°09′53″ S
,
146°51′01″ E
],
11 Feb. 2016
;
V. Ryland
leg.;
on
Ipomoea
pes-caprae;
RBINS
• 1 ♂,
1 ♀
; same collection data as for preceding; QM
•
1 ♀
;
Magnetic Island
,
Geoffrey Bay
Beach; [coordinates of
Geoffrey Bay
:
19°09′17″ S
,
146°51′54″ E
];
10 Feb. 2016
; V. Ryland leg.;
on
Ipomoea
pes-caprae; RBINS
• 1 ♂; same collection data as for preceding; QM
• 1 ♂, 1 ♀; same collection data as preceding;
24 Feb. 2016
; RBINS
• 2 ♂♂, 1 ♀; same collection data as for preceding; QM
• 1 ♂,
1 ♀
;
Magnetic Island
,
Picnic Bay
Beach; [coordinates of
Picnic Bay
:
19°10′54″ S
,
146°50′27″ E
];
5 Feb. 2016
; V. Ryland leg.;
on
Ipomoea
pes-caprae; RBINS
• 1 ♀; same collection data as for preceding; QM
• 1 ♀; same collection data as preceding;
12 Feb. 2016
; RBINS
• 1 ♀; same collection data as preceding;
18 Feb. 2016
; QM
• 1 ♀; same collection data as preceding;
27 Feb. 2016
; RBINS
• 1 ♂, 2 ♀♀; same collection data as for preceding; QM
• 1 ♂; same collection data as preceding;
10 Jan. 2016
;
on
Eucalyptus
sapling;
RBINS
• 1 ♂;
Magnetic Island
;
19°08′18″ S
,
146°50′04″ E
;
30 Mar. 1929
; QM
• 2 ♂♂,
1 ♀
;
Magnetic Island
;
A.M. Lea
leg.;
SAM
•
1 ♀
; same collection data as preceding;
G. F. Hill
leg.;
SAM
• 1 ♂,
3 ♀♀
;
Magnetic Island
;
8 Nov. 2006
;
J. Constant
leg.;
on
Canarium australianum
sapling;
RBINS
•
1 ♀
;
Magnetic Island
; 1914;
Dr Scharff
leg.;
BMNH
• 1 ♂;
Alligator Creek
;
19°23’56”S
146°55’36”E
;
1 Apr. 1934
; H. Hacker leg.; USNM
• 1 ♀; same collection data as preceding;
25 Apr. 1934
; USNM.
Description
MEASUREMENTS AND RATIOS. LT: ♂ (n = 9):
7.1 mm
(6.2–7.7);
♀
(n = 3):
7.7 mm
(7.5–8.0); BV/LV = 4.5; BF/LF = 1.68; LP+LM/BT = 0.68; Ltg/BTg = 2.35; LW/BW = 1.86.
Male
HEAD (
Figs 16A–D
,
17A–D
,
18A–D
). Vertex concave, with anterior and posterior margins parallel, curved; brown with darker marking at lateral angles. Frons uniformly black-brown. Clypeus elongate, entirely black-brown. Genae black-brown with yellowish markings along anterior margin. Labium black, reaching metacoxae. Antennae black-brown; scape short, ring-shaped; pedicel subcylindrical, slightly narrowing towards apex.
THORAX (
Figs 16A–D
,
17A–D
,
18A–D
). Pronotum brown with small yellowish spot on each side; slightly wrinkled; obsolete median carina and 2 small impressed points on disc. Lateral fields of prothorax dark brown. Mesonotum brown variegated with blackish; yellowish spot on middle of anterior margin and at apex of scutellum; median and peridiscal carinae weakly marked; median carina stopping before scutellum; slight impression before scutellum. Red ventrally. Tegulae dark brown.
TEGMINA (
Figs 16A, C
,
17A, C
,
18A, C
). Brown slightly variegated with yellowish and black; often pale yellowish marking on vein A1 at midlength of clavus; marked with black along costal margin, more broadly so on posterior half, and along posterior margin (
Figs 16A, C
,
17A, C
); sometimes a broad, whitish, transverse band at proximal ½, bordered with black markings along costal margin (
Fig. 18A, C
). Darker, median, irregular marking at apical ¾; triangular white marking on costal margin on nodal line, sometimes reduced; no white spot at apicosutural angle. Costal and sutural margins subparallel; costal margin slightly sinuate; apical margin obliquely rounded.
POSTERIOR WINGS (
Figs 16E
,
17E
,
18E
). Dark brown, paler on anal area and with large blackish area reaching apical margin; elongate, transverse, subtriangular white marking at apicocostal angle, extending on 3–4 cells. Margin of anal area slightly sinuate; sutural margin with 1 cleft, cubital one not marked.
LEGS (
Figs 16A–D
,
17A–D
,
18A–D
). Pro- and mesocoxae black-brown. Pro- and mesofemora blackbrown with reddish spots marking obsolete rings. Pro- and mesotibiae black-brown with reddish markings on 3 obsolete rings. Pro- and mesotarsi black-brown with basal half of third tarsomere paler. Metacoxae reddish; metafemora reddish with apex brown. Metatibiae brown, with 3 lateral spines paler basally and 8 apical black-brown spines. Metatarsi brown, with a ventral row of 6 black spines on first tarsomere.
ABDOMEN (
Fig. 18F
). Bright red with genital segments black-brown.
MALE GENITALIA (
Fig. 19
). Posterior margin of pygofer in lateral view strongly sinuate, strongly roundly projecting at dorsal ½, rather broad ventrally (
Fig. 19A, C
). Anal tube spatulate, 2.2 times as long as broad, laterally constricted at level of epiproct, slightly curved ventrally in lateral view; lateral margins slightly curved ventrally on apical ¾; apical margin rounded (
Fig. 19 A–B
). Gonostyli fused on basal third of length of centroventral part and projecting posteriorly (
Fig. 19A, C
). Centroventral part broad and dorsoventrally flattened on basal third, then strongly narrowing into a long spinose process strongly sinuate basally in lateral view and ending in narrow hook curved ventrally (
Fig. 19A, C
). Laterodorsal part of gonostyli strongly elongate and curved posteroventrally, surpassing level of centroventral part ventrally; lateral process elongate, projecting laterally and longer than spoon-shaped process (
Fig. 19A, C–D
). Dorsal portion of phallobase with hooked process on each side, progressively narrowing from base to apex, directed posterocentrally and with apex pointing dorsally (
Fig. 19E–F
). Ventral portion of phallobase trilobed in dorsal view, with median lobe surpassing phallus (
Fig. 19E–F
). Phallus dorsoventrally flattened, rather broad, with apical margin emarginate in middle (
Fig. 19E–F
).
Fig. 15.
Olonia marginata
Distant, 1906
, lectotype, ♀.
A
. Habitus, dorsal view.
B
. Habitus, ventral view.
C
. Habitus, lateral view.
D
. Habitus, normal view of frons.
E
. Labels.
Fig. 16.
Olonia marginata
Distant, 1906
, ♂ from mainland.
A
. Habitus, dorsal view.
B
. Habitus, ventral view.
C
. Habitus, lateral view.
D
. Habitus, normal view of frons.
E
. Posterior wing.
Fig. 17.
Olonia marginata
Distant, 1906
, dark form, ♂ from Magnetic Island.
A
. Habitus, dorsal view.
B
. Habitus, ventral view.
C
. Habitus, lateral view.
D
. Habitus, normal view of frons.
E
. Posterior wing.
Fig. 18.
Olonia marginata
Distant, 1906
, form with pale bands, ♂ from Magnetic Island.
A
. Habitus, dorsal view.
B
. Habitus, ventral view.
C
. Habitus, lateral view.
D
. Habitus, normal view of frons.
E
. Posterior wing.
F
. Abdomen, dorsal view.
Fig. 19.
Olonia marginata
Distant, 1906
, ♂, genitalia.
A
. Pygofer, anal tube and gonostyli, left lateral view.
B
. Anal tube, dorsal view.
C
. Pygofer and gonostyli, ventral view.
D
. Laterodorsal part of left gonostylus, dorsal view.
E
. Aedeagus, left lateral view.
F
. Aedeagus, dorsal view. Abbreviations:
An
= anal tube;
G
= gonostyli;
Py
= pygofer.
Fig. 20.
Olonia marginata
Distant, 1906
, ♀ from mainland.
A
. Habitus, dorsal view.
B
. Habitus, ventral view.
C
. Habitus, lateral view.
D
. Habitus, normal view of frons.
E
. Posterior wing.
F
. Abdomen, dorsal view.
Fig. 21.
Olonia marginata
Distant, 1906
, ♀ from Magnetic Island.
A
. Habitus, dorsal view.
B
. Habitus, ventral view.
C
. Habitus, lateral view.
D
. Habitus, normal view of frons.
E
. Posterior wing.
F
. Abdomen, dorsal view.
Female
Similar to male, but with white spot at apicosutural angle of posterior wing reduced, extending on 1 cell (
Figs 15
,
20
). Sometimes with a reduced white transverse band at basal ½ of tegmina visible in a marking along costal and sutural margins (
Fig. 21
).
Nymph
HEAD (
Fig. 22
). Brown, as broad as pronotum, frons flattened, with lateral rounded row of yellowish tubercles; group of yellowish tubercles dorsally on each side of joint between frons and vertex; clypeus reaching mesocoxae.
Fig. 22.
Olonia marginata
Distant, 1906
, nymphs from Townsville, Bushland Beach, 4 Apr. 2016.
A–E
. Fourth instar nymphs. A. Dorsal view. B. Ventral view. C–E. Lateral view.
F–G
. Fifth instar nymph. F. Lateral view. G. Dorsal view.
THORAX (
Fig. 22
). Brown; pronotum short, with two dense rows of yellowish tubercles on each side; meso- and metanotum with a circular group of yellowish tubercles at base of wing buds and some tubercles near apex of wing buds.
LEGS (
Fig. 22
). Brown slightly variegated with yellowish; large in comparison to body size. Anterior and median femora and tibiae broad and dorsoventrally flattened; posterior tibiae with three strong lateral spines.
ABDOMEN (
Fig. 22
). Rather short, less than half as long as head and thorax combined, and narrower than thorax; pale yellowish bown with dorsal median line and apex darker. Live specimens with abdomen covered in white wax, except three apical segments with brown wax; each segment bearing a waxy plate on each side; two movable, very elongate, waxy filaments at apex of abdomen (
Figs 23B–C
,
25B–C, K, N
).
Fig. 23.
Olonia marginata
Distant, 1906
, in nature, Townsville.
A–C
. Bushland Beach, 4 Apr. 2016. A. Habitat. B–C. Nymph on
Ipomoea
pes-caprae
(L.) R.Br.
D
. Egg case on
Ipomoea
pes-caprae
, Rowes Bay Beach, 25 Feb. 2016.
E–F
. Townsville Port Wall, 15 Mar. 2016. E. Habitat. F. Female on
Ipomoea
pes-caprae
.
G
. Female parasitized by larva of Epipyropidae (
Lepidoptera
), Rowes Bay Beach, 26 Feb. 2016.
H
. Habitat, Small Boat Harbour, 19 Feb. 2016.
I
. Habitat, Strand Beach, 15 Mar. 2016.
J
. Habitat, Pallarenda Beach, 25 Feb. 2016. Photographs by V. Ryland.
Distribution and biology
This species is recorded from an area around Townsville in North
Queensland
(
Fig. 3
), in the Einasleigh Upland Savanna,
Queensland
Tropical Rainforests and Brigalow Tropical Savanna bioregions. Adult specimens were collected in January, February, March, April, August, Sepember and October, while nymphs were observed in Feb. and Apr. (
Figs 23B–C
,
25B–C, K
), and an egg mass in Apr. (
Fig. 23D
), leading to the conclusion that the species is probably present most of the year. A caterpillar of an unidentified species of Epipyropidae (
Lepidoptera
) was observed on the back of a female specimen (
Fig. 23G
).
Fig. 24.
Olonia marginata
Distant, 1906
, in nature, Magnetic Island, 8 Nov. 2006, on
Canarium australianum
F.Muell.
saplings.
A–B
. Male on 2 m tall sapling.
C
. Two m tall sapling of
C. australianum
.
D–E
. Female on small
C. australianum
sapling.
F
. Terminal leaves of small
C. australianum
sapling.
G
. Leaves of
C. australianum
.
H
. Fruits of
C. australianum
.
I
. The author and his 5 month old daughter Emilie observing specimens of
O. marginata
. Photographs by the author except I by L. Despontin.
Fig. 25.
Olonia marginata
Distant, 1906
, in nature, Magnetic Island.
A–G
. Picnic Bay Beach. A. Habitat, 6Feb.2016.B–C.Nymph on
Ipomoea
pes-caprae
(L.) R.Br, 6Feb.2016.D.Adult on
Ipomoea
pes-caprae
, 6 Feb. 2016. E. Pair on
Corymbia tesselaris
K.D.Hill & L.A.S.Johnson
regrowth, 31 Jan. 2016. F. Male on
Eucalyptus
sp. sapling, 10 Jan. 2016. G. Female on
Pipturus argentaeus
Wedd.
, 29 Jun. 2014.
H–K
. Nelly Bay, 11 Feb. 2016. H. Habitat. I. Adults on
Ipomoea
pes-caprae
. J. Male. K. Nymph on
Ipomoea
pes-caprae
.
L–O
. Geoffrey Bay. L. Habitat, 10 Feb. 2016. M. Male on
Ipomoea
pes-caprae,
10 Feb. 2016. N. Nymph on
Ipomoea
pes-caprae,
10 Feb. 2016. O. Female on
Ipomoea
pes-caprae,
24 Feb. 2016. Photographs by V. Ryland.
Olonia marginata
is polyphagous and has been collected or observed on
Melaleuca quinquenervia
(Cav.) S.T.Blake (Myrtaceae)
,
Eucalyptus
sp. (
Myrtaceae
) (
Fig. 25F
),
Canarium australianum
F.Muell. (Bursaceae)
(
Fig. 24
),
Vitis vinifera
L. (
Vitaceae
),
Pipturus argentaeus
Wedd. (Urticaceae)
(
Fig. 25G
),
Corymbia tesselaris
K.D.Hill & L.A.S.Johnson (Myrtaceae)
(
Fig. 25E
) and in numbers on
Ipomoea pescaprae
(L.) R.Br. (
Convolvulaceae
) (
Figs 23B–D, F
,
25B–D, I–K, M–O
), on stems and young branches with a thin skin.
It is interesting to note that the 2016 sampled locations in Magnetic Island and Townsville (
Figs 23A, E, H–J
,
25A, H, L
) were all more or less destroyed by the Tropical Cyclone Yasi in
Feb. 2011
, which covered these habitats in smashing waves and left them buried under sand and beach debris (pers. com. V. Ryland,
Feb. 2016
). The species has, however, re-colonised the areas in less than five years, displaying a good degree of adaptation to habitat disturbance.