The henicopid centipede Haasiella (Chilopoda: Lithobiomorpha): new species from Australia, with a morphology-based phylogeny of Henicopidae
Author
Edgecombe, Gregory D.
text
Journal of Natural History
2004
2004-01-10
38
1
37
76
http://www.tandfonline.com/doi/abs/10.1080/0022293021000007552
journal article
10.1080/0022293021000007552
1464-5262
4653649
3BCB0BE6-781E-46ED-9BA4-FF79E5644F35
Paralamyctes
(
Haasiella) cammooensis
n. sp.
(figures 2–5, 6A–D, 7–12, 38B)
Etymology.
From Cammoo Caves, the
type
locality.
Diagnosis.
Member of
P
.
(
H
.)
cammooensis
Group with four or five teeth on dental margin of maxillipede; legs finely setose, without stronger, spine-like setae on prefemur and femur; few setae on first genital sternite of female.
Holotype
.
ANIC-03-1, female (figure 2),
Cammoo Caves
(now
Mt Etna Caves
NP
), near
Rockhampton
,
Queensland
, 23°10∞S, 150°28∞E,
R
.
W. Taylor
and
T
.
A. Weir
,
25 October 1976
, dense, low closed forest (ex
ANIC
Berlesate 535). Length of head 0.56 mm; length of body 5.6 mm.
Paratypes
.
figured specimens ANIC-03-2–ANIC-03-4, ANIC-03-7, four females, ANIC-03-5, ANIC-03-6, two males; unfigured specimens ANIC-03-8 (five females, nine males); all from type locality, same collection.
Other material
.
NE
Queensland
rainforest: ANIC-03-9, male,
Moses Creek
,
4 km
N by
E Mt Finnigan
, 15°47∞S, 145°17∞E,
T
.
Weir
,
14–16 October 1980
;
ANIC-03-10, female,
Mt Lewis
,
2 mi
above Bushy Creek
, 16°35∞S, 145°18∞
E, J. G. Brooks
,
25 November 1969
;
ANIC-03-11, male,
Mt Lewis
, 16°35∞S, 145°13∞E,
1010 m
,
R
. W.
Taylor
and J.
Feehan
,
20 June 1971
;
ANIC-03-12, three females, five males, Black
Mt Rd
,
30 km
N Kuranda
, 16°38∞S, 145°30∞
E, J. G. Brooks
,
4 November 1969
;
ANIC-03-13, female, Black
Mt Rd
,
15 mi
N Kuranda
, 16°42∞S, 145°31∞E, A. Walford-
Huggins
,
6 November 1975
;
ANIC-03-14, three females, four males, six juveniles, larva
IV
,
Black Mt Rd
, 13.3 km N
Kuranda
, 16°45∞S, 145°34∞E,
ca
1500 ft
,
J. G. Brooks
,
3 January 1971
;
ANIC-03-15, male,
13 km
NW Kuranda
, 16°45∞S, 145°34∞E, J.
Doyen
,
6 December 1982
;
ANIC-03-16, two males, Black
Mt Rd
, 16°45∞S, 145°35∞E,
430 m
,
R
.
W. Taylor
and
J. Feehan
,
27 June 1971
;
ANIC-03-17, two females, Black
Mt Rd
,
7 km
NNW Kuranda
, 16°45∞S, 145°37∞E,
400 m
,
A. Calder
and
T
.
Weir
,
30 March 1984
;
ANIC-03-18, male, Black
Mt Rd
, 11.2 km N
Kuranda
, 16°46∞S, 145°35∞
E, J. G. Brooks
,
30 April 1970
;
ANIC-03-19, female,
1 km
W Kuranda
, 16°50∞S, 145°37∞E, J.
Doyen
,
6 December 1982
;
ANIC-03-20, female,
Crystal Cascades
, 16°57∞S, 145°42∞
E, J. G. Brooks
,
30 April 1970
;
ANIC-03-21, female, male,
Mt Tiptree
, 17°03∞S, 145°38∞E,
730 m
,
R
. W.
Taylor
and J.
Feehan
,
29 June 1971
;
ANIC-03-22, two females, male, near
Mt Haig
, 17°07∞S, 145°35∞E,
1000 m
,
R
. W.
Taylor
and J.
Feehan
,
30 June 1971
;
ANIC-03-23, male,
Boar Pocket Rd
,
5 mi
N Gillies Hwy
, 17°10∞S, 145°39∞
E, J. G. Brooks
,
9 December 1969
;
ANIC-03-24, female,
Gillies Hwy
,
2 km
E Mt Nomico
, 17°13∞S, 145°42∞E,
720 m
,
R
.
W. Taylor
and
T
. A.
Weir
,
28 October 1976
;
ANIC-03-25, male,
Mt Bellenden Ker
, summit, 17°16∞S, 145°51∞E,
1560 m
,
R
. W.
Taylor
and J.
Feehan
,
7 July 1971
;
ANIC-03-26, male,
27 km
S Atherton
,
Longlands Gap Rd
, 17°28∞S, 145°29∞
E, J. G. Brooks
,
11 November 1969
;
ANIC-03-27, two males, two larvae
IV
,
Crawford’s Lookout
, 17°37∞S, 145°48∞E,
320 m
,
R
. W.
Taylor
and J.
Feehan
,
5 July 1971
;
ANIC-03-28, four females, three males, 1.6 km SE
Crawford’s Lookout
, 17°37∞S, 145°48∞E,
ca
300 m
,
R
.
W. Taylor
and
J. Feehan
,
10 July 1971
;
ANIC-03-29, two females, male,
McNamee Creek
, 17°40∞S, 145°48∞E,
ca
400 m
,
R
. W.
Taylor
and J.
Feehan
,
8 July 1971
;
ANIC-03-30, two juveniles,
1 km
N Tully Falls
NP, 17°45∞S, 145°32∞E, A.
Walford-Huggins
,
8 January 1976
;
ANIC-03-31, female,
Lacey’s Creek
, 17°52∞S, 146°04∞E,
ca
40 m
,
R
. W.
Taylor
and J.
Feehan
,
9 July 1971
;
ANIC-03-32, two females, four males, 2.5 mi W
Paluma
, 19°01∞S, 146°11∞E,
2940 ft
,
J. G. Brooks
,
13 January 1974
;
ANIC-03-33, two females, 2.7 mi W
Paluma
, 19°01∞S, 146°11∞E,
2920 ft
,
J. G. Brooks
,
11 January 1975
;
AM
KS 74958
, male,
Danbulla
SF,
Robson Creek Rd
, 17°07∞S, 145°37∞
E, G. Milledge
,
27 April to 2 May 1998
;
QM
S.45157, male,
Bellenden Ker Range
, summit
TV
station, 17°16∞S, 145°51∞E,
1560 m
,
Earthwatch
/
Qld. Museum
,
17 October to 7 November 1981
.
FIGS 2, 3.
Paralamyctes
(
Haasiella
)
cammooensis
n. sp.
(2) Holotype, ANIC-03-1, female, Cammoo Caves, Mt Etna Caves NP, Queensland, scale 0.5 mm. (3) AM KS 74958, male, Danbulla SF, Queensland, scale 0.5 mm.
FIG. 4.
Paralamyctes
(
Haasiella
)
cammooensis
n. sp.
Scanning electron micrographs. All from Cammoo Caves, Mt Etna Caves NP, Queensland. (A–D) ANIC-03-5, male. (A, B) Dorsal and ventral views of head, scales 100
µ
m; (C) telopodite and coxosternite of maxillipede, scale 50
µ
m; (D) dental margin of maxillipede coxosternite, scale 10
µ
m. (E–G) ANIC-03-3, female; (E, F) maxillipede and detail of dental margin of coxosternite, scales 50
µ
m, 10
µ
m; (G) first maxillae, scale 20
µ
m. (H) ANIC-03-2, female, coxal processes and telopods of first maxillae, scale 20
µ
m.
NSW: ANIC-03-34, female, male, Barrington Tops SF, Dilgry River, 31°53∞S, 151°32∞E,
T
. Weir and
A. Calder
,
15–16 November 1981
, open forest
;
AM
KS 67183
, female,
Kanangra-Boyd NP
,
Blood Filly Creek
, 33°51∞S, 150°03∞
E, M. Gray, G
. Hunt and
J. McDougall
,
27 March 1976
,
Eucalyptus fastigata
-viminalis
.
Description
Length up to 7.1 mm. Head shield of subequal length and width (length 0.82 mm, width 0.80 mm in largest specimen). Head shield usually light brown; purple-black pigment concentrated around ocellus; antenna pale orange or yellow; tergites typically light brown; fresher specimens yellow to pale yellow-brown with purple-black mottling concentrated in median longitudinal band and on borders; legs pale brown; sternites occasionally pale lavender, with similar pigmentation on legs except for dull yellow tarsi.
Head shield smooth. Median notch at anterior margin moderately deep; posterior margin of head transverse or weakly convex. Median furrow well impressed to transverse suture (figure 4A), with indistinct, weak to strong continuation behind transverse suture to middle of head shield. Antenna extending back to TT5 (figure 2) or 6, usually with 17 articles, exceptionally 16, other combinations include 14
/
18, 17
/
19, 17
/
20, 18
/
18 articles; basal two articles moderately larger than succeeding articles; articles 3–16 variable in length, a few not much longer than wide; articles gently widening distally; distal article typically about 1.7 times length of penultimate article. Typical henicopid increase in abundance of trichoid sensilla from segment 3, dense from segment 5 or 6; most trichoid sensilla orientated anteriorly (figure 6D); articles 3–16 bearing a club-shaped thin-walled basiconic sensillum at anterior edge beside band of arthrodial membrane (figure 5A). Ocellus large, pale, with faint independent convexity. Tömösváry organ large, elliptical, mid-length at about 35% length of cephalic pleurite; outer margin slightly inward of lateral edge of pleurite (figure 6A).
FIG. 5.
Paralamyctes
(
Haasiella
)
cammooensis
n. sp.
Scanning electron micrographs. (A, I) ANIC-03-2, female, Cammoo Caves, Mt Etna Caves NP, Queensland. (A) Third- and fourth-last antennal articles, scale 5
µ
m; (I) gonopod, scale 10
µ
m. (B–G) QM
S45157
, male, Bellenden Ker Range, Queensland. (B) Mandible, scale 50
µ
m; (C) inner (posterior) view of mandibular gnathal lobe, scale 10
µ
m; (D) aciculae and fringe of branching bristles, scale 10
µ
m; (E) mandibular teeth, scale 10
µ
m; (F) furry pad and accessory denticles, scale 5
µ
m; (G) aciculae, scale 5
µ
m. (H) ANIC-03-3, female, locality as for A, tarsus and pretarsus of second maxilla, scale 15
µ
m.
FIG. 6. (A–D)
Paralamyctes
(
Haasiella
)
cammooensis
n. sp.
Cammoo Caves, Mt Etna Caves NP, Queensland. (A) ANIC-03-3, female, cephalic pleurite, showing Tömösváry organ, scale 20
µ
m. (B, C) ANIC-03-5, male, gonopod and detail of terminal process, scales 10
µ
m, 5
µ
m. (D) ANIC-03-7, female, articles in distal part of antenna, scale 50
µ
m. (E–H)
Paralamyctes
(
Haasiella
)
subicolus
n. sp.
(E, F) QVMAG 23:5282, female, Christmas Hills, Tasmania. (E) Coxa of second maxilla, scale 100
µ
m; (F) cephalic pleurite, showing Tömösváry organ, scale 50
µ
m. (G, H) QVMAG 23:22969, male, Wombat Hill, Waratah, Tasmania, gonopod and detail of terminal process, scales 30
µ
m, 10
µ
m.
FIGS 7–12.
Paralamyctes
(
Haasiella
)
cammooensis
n. sp.
(7–10) ANIC-03-32, male, 2.5 mi W Paluma, Queensland, scales 0.5 mm. (7) Leg 12. (8) Leg 13. (9) Leg 14. (10) leg 15. (11) ANIC-03-6, male, Cammoo Caves, Mt Etna Caves NP, Queensland. Terminal segments and gonopods, scale 0.25 mm. (12) ANIC-03-4, female, Cammoo Caves. Terminal segments and gonopods, scale as for figure 11.
Clypeus with row of four setae immediately in front of labrum; cluster of five setae at anteromedial apex includes two pairs along margin and single medial seta. Transverse seta projects medially from labral sidepiece; labral margin weakly curved along inner extent of sidepiece to where cluster of branching bristles overhangs; bristles branching along their entire length.
Maxillipede
. Most of lateral margin of coxosternite strongly converging forwards, convergence less in anterior portion including dental margin (figure 4C); each half of dental margin sloping backwards medially, most commonly with 4+4 teeth (figure 4F), frequently with 4+5, 5+4 or 5+5 (figure 4C), one specimen with 6+6; teeth conical, pointed, gradually diminishing in size medially; porodont or pseudoporodont absent; median notch variably lacking (figure 4D) or present, V-shaped (figure 4F); setae sparsely scattered on coxosternite. Pretarsal part of forcipular tarsungulum about equal in length to tarsal part (figure 4C). Setae evenly scattered on tarsal part of tarsungulum, tibia and femur.
Mandible
. Trunk with single fissure defining laminia condylifera (figure 5B). Four paired teeth (figure 5C). Five plumose aciculae; pinnules more or less symmetrical on dorsal and ventral margins of aciculae, most digitiform, with blunt tips, maximum of 9 or 10 pinnules along each side of acicula (figure 5G), as few as three along some aciculae in some specimens. Fringe of branching bristles skirts aciculae (figure 5D), shortening dorsally to narrow fringe beneath furry pad (figure 5E, F); bristles narrow-based, regularly branching along entire length, including bases (figure 5D). Three dorsal teeth bearing blunt triangular to rather scale-like accessory denticles; field of accessory denticles continuous between teeth; outer row of denticles on each tooth delineated by strong groove
/
grooved ridge (figure 5E). Furry pad separated from accessory denticle field on teeth by band of large, flattened scales (figure 5F); bristles of furry pad relatively short, most markedly tapering, with simple, pointed tips, occasionally bifurcate at tip.
First maxilla
(figure 4G, H). As described below for
P
.
(
H
.)
subicolus
except coxal process narrow, tapering, bearing three simple setae at tip, one seta medially near its base; simple setae on distal article of telopod mostly confined to membranous strip near inner margin, only a few tiny setae on more strongly sclerotized outer two-thirds of article.
Second maxilla
(figure 5H). As described below for
P
.
(
H
.)
subicolus
except fewer (four or five) short setae in band across anterior part of coxa.
Tergites
. Tergites smooth. T1 trapeziform, 80–85% width of head, posterior angles rounded, posterior margin transverse or faintly convex backwards (figures 2, 3); T3 about 90% width of head, lateral borders subparallel; posterior angles of TT3 and 5 rounded, posterior borders faintly concave; TT1 and 3 bordered posteriorly, border on T5 variably complete; lateral borders of TT5–14 convex; T7 relatively short, posterior margin varying from weakly concave to weakly convex, posterior angle blunt to rounded; TT7–14 bordered laterally only; posterior margins of TT8–13 nearly transverse, at most weakly concave, posterior angles rounded, with no projections; posterior margin of T14 transverse to gently concave, without consistent sexual dimorphism, with blunt posterior angle; tergite of first genital segment and telson well sclerotized in both sexes. A few setae along lateral margins of tergites, longer setae anteriorly on long tergites, at posterolateral corner of short tergites; TT2, 4 and 9–14 with a few setae along posterior margin.
All setae on legs fine, none thickened or strongly pigmented, tarsal setae more slender than those on prefemur, femur and tibia; on anterior 13 legs setae on inner margin of tarsus sloping distally along distal two-thirds. Distal spinose projections on tibia of legs 1–12 (figure 7), absent on legs 13–15 (figures 8–10). Tarsal articulation undefined on legs 1–12, weak to moderately strong but unflexed on leg 13, strong and flexed on legs 14 and 15; basitarsus only slightly longer than distitarsus on legs 13–15. Pretarsal accessory claws as much as 50% length of main claw; short, distally directed ventral sensory spur beneath main claw (figure 38B).
Coxal pores
. On legs 12–15, round; almost invariably two pores on each leg in females, inner pore slightly smaller, one specimen with three pores on leg 15; usually one pore on each leg in males, exceptionally two on legs 14 and
15 in
larger specimens.
Female
(figure 12). First genital sternite sparsely setose, in some specimens bearing only a few setae along posterior part; posterior margin concave between condylic articulations of gonopods, gently curved backwards in median part. Basal article of gonopod bearing three to seven setae; two small, conical spurs; inner spur slightly smaller; bases of two spurs nearly side by side (figure 5I). Second article of gonopod bearing three or four fairly long setae; third article bearing one or two setae. Claw simple (figure 5I).
Male
(figure 11). Sternite of segment 15 with transverse posteromedian margin. Sternite of first genital segment undivided, bearing a few setae that describe a shallow V across posteromedian part; posteromedian margin of sternite bulging backwards between gonopods. Gonopod of three articles and terminal process; first article bearing row of three setae at mid-length; second article bearing one or two setae just behind mid-length; third article bearing one to three setae near mid-length (figure 6B); length of terminal process as much as combined length of second and third articles of gonopod; proximal half of terminal process weakly tapering, about as long as third article of gonopod, bearing several short spines distally (figure 6C), distal half a slender flagellum. Penis small, conical.
Discussion.
A few specimens from the Barrington Tops and Blue Mountains,
New South Wales
, are provisionally assigned to this species, all other material of which is from the Rockhampton region,
Queensland
, and Wet Tropics of northeastern
Queensland
. The
New South Wales
specimens conform to
Queensland
material of
P
.
(
H
.)
cammooensis
in having 4+4 maxillipede teeth, one pore on coxae
12–15 in
the male, and two pores on these coxae in females. The samples differ only in a relatively thicker tarsus on the anal leg in the one
New South Wales
specimen (a female) that retains that leg.