Heissophila macrotheleae, a new genus and new species of Plokiophilidae from Thailand (Hemiptera, Heteroptera), with comments on the family diagnosis

2006·1.2 MB·English

by  Schuh, Randall T

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Heissophila macrotheleae, a new genus
and new species of Plokiophilidae from
Thailand (Hemiptera, Heteroptera), with
comments on the family diagnosis1
R.T. SCHUH
Abstract: Heissophilamacrotheleaenov.gen. et nov.sp., is described from the webs of MacrotheleAUSSE-
RER sp. (Araneae: Mygalomorphae: Hexathelidae) in Thailand. Digital macrophotographs, scanning
electron micrographs, and line drawings are provided for a range of morphological structures. The ta-
xon is distinguished from previously described arachnophilic Plokiophilidae by the 3-segmented tarsi,
found elsewhere only in the New World taxon Lipokophila SˇTYS, and the apparent absence of traumatic
insemination as evidenced from the structure of the male genitalia which lack the piercing acus found
in all other described plokiophilid taxa.
Key words: classification, Heteroptera, Macrothele, new genus, Plokiophilidae.
Introduction clypeus-costal fracture 1.27 mm, width
pronotum 0.68 mm. Coloration(Fig. 1A, B):
The Plokiophilidae are the subject of few
Castaneous. Surface and Vestiture(Figs 1A,
works, the only comprehensive treatment
B, E, 2A, B): Vestiture of dorsum and venter
being that of CARAYON (1974). This no
comprising short, reclining, simple setae.
doubt derives from the fact that the group is
Antennae and tibiae with reclining setae of
seldom encountered by general collectors
uniform length. Head probably with 4 pairs
owing to their obscure habit of living only in
of long, suberect cephalic setae, two pairs sit-
the webs of a few spiders and of the Em-
uated near level of antennal insertion, one
bioptera. Material on which the present pa-
pair lateromedially on frons, and posterior
per is based was generously provided to the
pair laterad of ocelli (Fig. 2B); pronotum ap-
author by Ernst Heiss, Innsbruck, Austria.
parently without macrocheta on anterolater-
Heissophilanov.gen. (Figs 1-5) al angle as recorded by CARAYON(1974) for
other Plokiophilidae; abdominal segment 8
Type species: Heissophila macrotheleae, new
with a macrocheta laterally (Fig. 4A).
species
Diagnosis: Recognized uniquely among
Structure: Head (Figs 1A, B, 2A, B):
Old World Plokiophilidae by the 3-segment-
Head roughly cylindrical, elongate antero-
ed tarsi, the short broad pygophore, the
posteriorly, projecting beyond anterior mar-
short, stout, right-angled parameres, the four
gin of eyes by about 1.5 time length of eye;
veins (dead) in the membrane, the endoso-
vertex and frons sloping at same angle to
ma in repose in the form of a tube with many
nearly vertical and weakly prominent
internal spicules, and the presence of ovipos-
clypeus; buccal cavity more or less round, di-
itor valves within the female abdomen.
rected anteroventrally, greatly removed
Description: Male: Small, rather stout from posterior margin of head by gula of
bodied; total length 2.12 mm, length apex length greater than diameter of eye (Fig.
1The taxon is named in honor of Ernst Heiss, on the occasion of his seventieth birthday, in recognition of his con-
Denisia 19, zugleich Kataloge
tributions and dedication to heteropterology, particularly the Aradidae, as well as his generosity and friendship over der OÖ. Landesmuseen
the years to all of us interested in the study of the true bugs. Neue Serie 50 (2006), 637–645
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Fig. 1: Heissophila
macrotheleae. (A)
Male habitus. (B)
Female habitus. (C)
Membrane, light
micrograph. (D)
Corium and
membrane, scanning
electron micrograph.
(E) Corial process,
dorsal view, scanning
electron micrograph.
(F) Corial process,
ventral view, scanning
electron micrograph
(cf, costal fracture; cs,
campaniform
sensillum; pc, corial
process).
2A); mandibular and maxillary plates small; removed from anterior margin of pronotum
buccula very narrow (Fig. 2A); labium long, by about the diameter of antennal segment
slender, tapering to acute apex, slightly sur- 1; eyes in lateral view occupying about half
passing apex of middle coxae, segment 1 of height of head, nearly semicircular, poste-
short and broad, well developed (Figs 2A, rior margin nearly straight; antennal fossa
4H), ratio of length of segments 1-4, 2:4:4:7 located just below midpoint of eye and
(Fig. 4H); eyes relatively small in dorsal slightly removed from eye (Fig. 2A); ocelli
view, broadly excavated on mesial margin, small (Figs 1A, B, 2A, B), widely separated,
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Fig. 2: Heissophila macrotheleae, scanning micrographs of morphological structures. (A) Lateral view of head, male. (B) Dorsolateral
view of head, female, arrows indicating cephalic setae. (C) Compound eye, third (?) instar nymph. (D) Lateral view of tarsus, foreleg,
arrows indicating segmental divisions. (E) Lateral view of pretarsus, fore leg. (F) Frontoventral view of pretarsus, fore leg. (G) Detail of
ventral spines on segment 3 of fore leg tarsus (mc, macrocheta).
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placed at level of posterior margin of eye, re- moderately long, 3 segmented, segment 1
moved from eye by distance slightly greater very short, segments 2 and 3 subequal in
than diameter of ocellus (Fig. 2B); antenna length (Figs 2D, 4J, K); fore and middle tar-
relatively short, segment 1 short, not attain- si with a pair of strong reclining spinelike se-
ing apex of head (Fig. 2A), segments 2, 3, tae on segment 3, removed from apex of tar-
and 4 subequal in length, segment 1 of sus by about 2 times length of spine (Figs
slightly greater diameter than segment 2, 2D, G, 4J); claws elongate, very slender,
nearly 2 times diameter of segments 3 and 4, roughly cylindrical, tapering to sharply
prepedicellite present as a narrow sclero- pointed apex, inner claw longer than exteri-
tized band at base of segment 2. Thorax: or (Figs 2D-F, 4J, K); parempodia well de-
Pronotum trapezoidal, lateral margins near- veloped, setiform, of unequal length (Fig.
ly straight, posterior margin excavated me- 2E, F), of similar structure on all 3 pairs of
dially and convexly rounded laterally; legs, longer parempodium on side of shorter
pronotum strongly elevated posteriorly, claw. Abdomen: (Figs 3A, 4A): Short, stout;
transversely rounded (Fig. 1A, B); calli sterna entire, ventrolateral tergites fused
small, ovoid, shining, and devoid of setae, with sterna but demarcated by a sublateral
widely separated and placed near lateral impression (Fig. 3A); terga widely separated
margins of pronotum (Fig. 1A, B); scutellum from dorsal laterotergites by a broad mem-
slightly wider than long. Metathoracic branous area, allowing for great expansion,
scent-gland evaporatory area of limited ex- as shown in Figure 3A for alcohol-perserved
tent, located anteroventrally on metepime- specimens; paired openings of dorsal abdom-
ron (Fig. 3D, inset), this structure noted as inal glands (DAG 1) visible on tergum 4
not discernible in Lipokophila by SˇTYS (Fig. 4A; but see female); spiracles placed
(1967). Hemelytra: Costal margin of cori- on sternum near lateral margin of abdomi-
um strongly convex, coriomembranal junc- nal segments 2-7. Genitalia (Figs 3A, B,
ture nearly straight, well defined, and weak- 4A-D): Pygophore: Short, very broadly
ly angled anteromedially (Fig. 1A-D); costal connected to abdomen, in contrast to gen-
fracture located at about two-thirds distance era such as LipokophilaSˇTYS; opening of py-
from base to apex of corium, angled antero- gophore directed dorsally (Figs 3A, 4A, C);
parameres arising laterally from a semicircu-
medially, traversing costal-radial veins, but
lar excavation with the dorsal thumblike
not beyond, lacking incisure on corial mar-
projection (Figs 3B, 4A, C, D). Aedeagus
gin (Fig. 1B); membrane with 4 longitudinal
(Fig. 4B, C): In repose endosoma enclosed
“dead“ veins, none of them bearing setae,
within phallosoma; phallobase with sclero-
the posterior 3 nearly straight, the most an-
tized arms directed anteriorly from attach-
terior rather strongly curving (Fig. 1A-C);
ment to capitate processes, broad basally, at-
corial process (processus corial of CARAYON
tenuated distally; anterodorsal portion of
1974) present sublaterally on membrane at
aedeagus with a “stirrup-like“ structure; en-
juncture of corium and membrane (Fig. 1D),
dosoma tubular, oriented in longitudinal ax-
dorsal surface bearing a single simple seta
is of body, anterior half appearing striated,
and apparently one campaniform sensillum
posterior half with many fine spicules di-
(Fig. 1E), ventral surface of corial process
rected internally and a pair of larger medial
bearing 4 campaniform sensilla (Fig. 1F).
spines directed posteriorly in repose (Fig.
Hind wing as in Figure 4; hamus absent. Co-
4B, C). Parameres (Fig. 4A, C, D): Sym-
rial Glands: Corial glands widely distrib-
metrical, narrow at base, body of paramere
uted, including at least pronotum, hemely-
broad and flattened, apical portion at nearly
tra, and antennal segment 2; external com-
right angle to body with a few short setae on
ponent in the form of an ovoid depression
apicodorsal margin.
with an elongate central mound, and
minute pore located at one end of mound Female (Fig. 1B): Structure and col-
(Fig. 3E-G). Legs: Relatively short; femora oration as in male; total length 2.09 mm,
of moderate length, slender, nearly parallel length apex clypeus-cuneal fracture 1.36
sided, devoid of spines; fore tibia with a mm, width pronotum 0.74 mm. Abdomen:
cleaning comb on medial surface at apex; no Paired openings of dorsal abdominal glands
fossula spongiosa; tarsi relatively stout, only visible on abdominal terga 4 and 5; spiracle
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Fig. 3: Heissophila macrotheleae, scanning micrographs of morphological structures. (A) Lateral view of abdomen, male. (B)
Lateral view of pygophore, male, detail. (C) Lateral view of apex of abdomen, female. (D) Lateral view of thoracic pleuron,
inset showing detail of scent-gland evaporatory area. (E) Antennal segment 2, arrow indicating integumental gland.
(F) Hemelytral surface, showing integumental gland. (G) Detail of integumental gland on hemelytron, arrow indicating pore
(mpe, metepimeron; mes, metepisternum; sce, scent-gland evaporatorium; sp 3, spiracle 3; sp 8, spiracle 8).
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Fig. 4:Heissophila
macrotheleae,
morphological
details.
(A) Male abdomen,
dorsal view.
(B) Aedeagus (top
to bottom), lateral,
dorsal, and
ventrolateral views.
(C) Pygophore and
aedeagus, cutaway
dorsal view.
(D) Left and right
parameres.
(E) Female
abdomen, dorsal
view.
(F) Female
abdomen, dorsal
cutaway view.
(G) Female
abdomen, ventral
cutaway view.
(H) Labium, lateral
view.
(I) Hind wing.
(J) Fore tarsus,
right leg, lateral
view.
(K) Hind tarsus, left
leg, lateral view.
(1A, first anal vein;
cu, cubital vein;
DAG 1, DAG 2,
opening of dorsal
abdominal gland;
m, medial vein; mc,
macrocheta; r,
radial vein; r+m,
radiomedial vein;
sp 8, spiracle 8; vlf,
valifer; vlv, valvula).
of segment 8 in female dorsal (Figs 3C, 4E). and tarsi pale. Compound eye with 7 or 8
Genitalia(Figs 3C, 4F, G): Ovipositor with facets and 2 central setae (Fig. 2C), similar
an elongate apparent valifer and short to condition seen in Lipokophila (SCHUH
valvulae (Fig. 4F, G); abdomen (macerated) 1993). Paired openings of dorsal abdominal
internally with a large baglike structure with
glands located between terga 3-4, 4-5, and
bulging bodies posterolaterally.
5-6, although these not visible with scan-
Nymph: General coloration red, tibiae ning electron microscopy.
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Etymology: A combination of Heiss and Tab. 1: Known associations of Plokiophilidae with spiders.
the suffix -phila, following pattern of gener-
Plokiophilidae Spider host Spider suborder and family
ic names previously proposed in the Plokio-
Heissophila Macrothele Mygalomorphae: Hexathelidae
philidae. Feminine.
Lipokophila Tengella Araneomorphae: Tengellidae (SCHUH1993)
Discussion: CARAYON (1961), in a brief Plokiophiloides Agelena Araneomorpha: Agelenidae (CARAYON1974)
article dealing with the distribution of vi- Plokiophiloides Ischnothele Mygalomorphae: Dipluridae (CARAYON1974)
viparity in the Heteroptera, recognized the
Plokiophilidae as a family distinct from the tum as occurring in all Plokiophilidae and as
Microphysidae, where its members had orig- being distinctively oriented in the two rec-
inally be placed by CHINA and MYERS ognized subfamilies. These setae are not ev-
(1929) and CHINA(1953). CARAYON(1961) ident in Heissophilanov.gen.
recognized two subfamilies, on the basis of
Heissophilanov.gen. is clearly a member
their being found in the webs of spiders and
of the Plokiophilidae, based on the presence
embiids, respectively, as well as the form of
of corial glands, head structure, and claw
viviparity, but gave no further diagnostic
structure. Nonetheless, other aspects of its
characters. SˇTYS (1967), in his description morphology are distinctly at variance with
and discussion of LipokophilaSˇTYS,explicitly those previously provided for most of the
deferred from more detailed morphological
Plokiophilidae. Heissophila nov.gen. and
characterization of CARAYON’s proposed Lipokophila SˇTYS are set apart from other
subfamilies. Later, CARAYON (1974) pro- plokiophilid genera by their possession of 3-
duced a comprehensive treatment of the
segmented tarsi (Figs 2D, 4J, K).
Plokiophilidae, including a more detailed
listing of diagnostic morphological features The pygophore of Heissophilanov.gen. is
for the family as well as of the two subfami- short and rather broad, as opposed to elon-
lies he had previously proposed. At the fam- gate and tubular, the condition seen in all
ily level, CARAYON (1974) concluded em- other known taxa. Possibly most notable is
phatically that the sclerotized tubular the structure of the aedeagus in the male.
processus gonopori (acus) is distinctive to The endosoma does not possess a processus
the Plokiophilidae, occurring in all known gonopori, but rather appears saclike in form,
members, that this structure is associated the distal half being covered with many, fine,
with the practice of traumatic insemination, closely-placed spicules, these being inwardly
and that consequently traumatic insemina- directed when the endosoma is in repose.
tion is a diagnostic feature for the group. Furthermore, the parameres in Heissophila
CARAYON(1974) for the first time provided nov.gen. are short, stout, and form a right-
a morphological characterization of the “co- angle bend subapically, rather than being
rial glands“, also treating them as diagnostic long, slender, and nearly straight as in all
for the group. Whereas CARAYON(1974) be- other known members of the Plokiophilidae.
lieved the corial glands to be restricted to
Examination of the female abdomen, in-
the external margin of the corium, in Heis-
cluding clearing and staining with chlorazol
sophilanov.gen. at least their external mani-
black, provided no indication of copulatory
festation is present on the coriaceous por-
tubes or other evidence of traumatic insem-
tion of the hemelytra, on the pronotum, and
ination. This would seem to be concordant
on antennal segment 2 (Fig. 3E-G). The
with the structure of the male phallus. Also,
structure of the head would also appear to be
the female has some structural elements as-
novel within the group, with its cylindrical
sociated with an ovipositor (Fig. 4F, G), in
form and relatively small eyes. Furthermore,
apparent contrast with other members of
all taxa currently placed in the Plokiophili-
the Plokiophilidae. I was not able to deter-
dae on the presence of corial glands and
mine whether Heissophila nov.gen. has a
head structure also have elongate asymmet-
spermatheca of the type described by
rically developed claws, and this might
CARAYON(1974) for some other members of
therefore be considered a diagnostic feature
the Plokiophilidae.
(see CARAYON1974; SCHUH1993). CARAY-
ON(1974) also listed the macrocheta locat- The hemelytral membrane in Heissophi-
ed on the anterolateral angle of the prono- la nov.gen. has four distinct longitudinal
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Fig. 5: Heissophila sus may be the basal condition in the group
macrotheleae,
rather than that the condition is uniquely
distribution,
derived in Lipokophila.
Thailand.
On the basis of some characters, Heis-
sophilanov.gen. would be placed in the sub-
family Plokiophilinae. These include the
overall structure of the legs and the propor-
tions of the scutellum. Yet, the distribution
of the corial glands is much wider in Heis-
sophilanov.gen. than is known for all other
Plokiophilidae, this in addition to features
of the male and female genitalia, which are
at variance with all other taxa placed with-
in the Plokiophilidae. For these reasons I am
treating the subfamily placement of Heis-
sophila nov.gen. as incertae sedis, pending
further work on morphology and relation-
ships within the Plokiophilidae.
Associations: Table 1 lists the presently
known associations of Plokiophilidae with
spiders. It appears that all spider taxa serv-
ing as hosts for Plokiophilidae construct
dead veins, contrary to the condition found funnel-shaped webs, even though several
in all other Plokiophilidae (see SˇTYS 1967) groups of distantly related spiders are in-
where the membrane appears to be devoid volved. It would also appear that most of
of veins, dead or alive, except for the corial these webs are large, persistent, and built by
process (noted by SˇTYS(1967); described in relatively large spiders.
detail by CARAYON (1974)). Nonetheless,
other members of the Cimicoidea such as Heissophila macrotheleaenov.sp.
some species of Anthocoridae possess 4 lon- (Figs 1-5)
gitudinal veins in the membrane. The struc- Diagnosis: See generic diagnosis
ture of the corial process is similar to that
Description: Measurements, male: To-
described for Plokiophiloides asolenCARAYON
tal length 2.12 mm, length clypeus-costal
(CARAYON1974), with several campaniform
fracture 1.27 mm, length head 0.20 mm,
sensilla, located primarily on the ventral
length pronotum 0.33 mm, length scutellum
surface of the wing (Fig. 1F).
0.27 mm, width head 0.31 mm, interocular
The costal fracture in Heissophila distance 0.22 mm, width pronotum 0.68
nov.gen. is readily seen with transmitted mm, width scutellum 0.31 mm, antennal
light, but difficult to see with scanning mi- segment 1-0.09 mm, 2-0.36 mm, 3-0.30 mm,
croscopy; there is no incisure on the costal 4-0.32 mm.
margin of the forewing. By contrast, evi-
Measurements, female: Total length
dence presented by CARAYON (1974) indi- 2.09 mm, length clypeus-costal fracture 1.36
cates that the costal fracture is at least read- mm, length head 0.20 mm, length pronotum
ily visible in most other species of Plokio- 0.35 mm, length scutellum 0.22 mm, width
philidae, while in Lipokophilait is prominent head 0.33 mm, interocular distance 0.21
and contributes to the formation of distinct mm, width pronotum 0.74 mm, width
cuneus (see SCHUH1993). scutellum 0.33 mm, antennal segment 1-
0.10 mm, 2-0.35 mm, 3-0.28 mm, 4-0.38
Thus, I tentatively propose that the di-
mm.
agnosis of the Plokiophilidae be altered to
include a modified description of the male Etymology: Named for its association
genitalia, possibly including the method of with the mygalomorph spider genus
insemination, and that the 3-segmented tar- Macrothele.
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Biotic Association: Heissophilamacrothe- Zusammenfassung
leae nov. sp. is known to occur only in the
Heissophila macrotheleae nov.gen. et
webs of Macrothele AUSSERER (Araneae:
nov.sp. wird aus Netzen der Spinnengattung
Hexathelidae).
MacrotheleAUSSERERsp. (Araneae, Mygalo-
Distribution: Chiang Rai and Sakon
morphae, Hexatelidae) aus Thailand neu
Nakhon Provinces, northern Thailand.
beschrieben. Morphologische Strukturen
Discussion: See generic discussion. Ex- werden durch Makrophotographien,
amined material is deposited in the Ameri- Rasterelektronenmikroskopische Photogra-
can Museum of Natural History (AMNH) phien und Strichzeichnungen dargestellt.
and the Museum d’Histoire Naturelle, Das Taxon unterscheidet sich von den bish-
Geneva. er beschriebenen arachnophilen Plokiophil-
idae durch 3-gliedrige Tarsen, die sonst nur
Holotype male: Thailand: Chang Rai Province:
Doi Tung, 20.32°N 99.82°E, 1300 m, 2 Oct 1992, in der Neuweltlichen Gattung Lipokophila
P.J. Schwendinger (AMNH_PBI 00137269) SˇTYSbekannt sind. Traumatische Insemina-
(AMNH). tion scheint nicht vorzuliegen, da die
Paratypes: Thailand: Chang Rai Province: Mae männlichen Genitalien – im Unterschied zu
Sai District: Doi Tung, 20.32°N 99.82°E, 1300
allen bisher bekannten Plokiophiliden –
m, 2 Oct 1992, P.J. Schwendinger, 9==
keinen sklerotisierten processus gonopori
(AMNH_PBI 00137271, AMNH_PBI
(acus) aufweisen.
00137315, AMNH_PBI 00139048), 4YY
(AMNH_PBI 00137270, AMNH_PBI
00137316, AMNH_PBI 00139048) (AMNH; References
MHNG); 30 Oct 1991, P.J. Schwendinger, 5==
(AMNH_PBI 00139046) (AMNH); 13 Oct CARAYONJ. (1961): La viviparite chez les Hétérop-
1994, P.J. Schwendinger, 2== (AMNH_PBI tères. — Verh. XI. Int. Kongr. Entomol. Wien
00139045), 2YY (AMNH_PBI 00139045) 1960 1: 711-714.
(AMNH). Sakon Nakhon Province: Kut Bak CARAYONJ. (1974): Etude sur les Hémiptères Plokio-
District: Phu Phan National Park, 16.82°N philidae. — Ann. Soc. Entomol. Fr. (N. S.) 10
104.43°E, 520 m, 25 Nov 1992, P.J. (3): 499-525.
Schwendinger, 3== (AMNH_PBI 00139044), CHINAW.E. & J.G. MYERS(1929): A reconsideration
1Y(AMNH_PBI 00139044) (AMNH). of the classification of the cimicoid families
Other Specimens Examined: Thailand: Chang (Heteroptera), with the description of two
Rai:Doi Tung, 20.32°N 99.82°E, 1300 m, 2 Oct new spider-web bugs. — Ann. Mag. Nat. Hist.
1992, P.J. Schwendinger, 3 nymphs (AMNH_PBI (10) 3: 97-125.
00139047, AMNH_PBI 00139048) (AMNH); CHINAW.E. (1953): A new subfamily of Microphysi-
30 Oct 1991, P.J. Schwendinger, 1 nymph dae (Hemiptera-Heteroptera). — Ann. Mag.
(AMNH_PBI 00139046) (AMNH). Sakon Nat. Hist. 12(6): 97-125.
Nakhon Province: Kut Bak District Co.: Phu
SCHUHR.T. (1993): Heteroptera: Family Plokiophili-
Phan National Park, 16.82°N 104.43°E, 520 m, dae. — In:EBERHARDW.G., PLATNICKN.I. & R.T.
25 Nov 1992, P.J. Schwendinger, 3 nymphs SCHUH(Eds),Natural history and systematics of
(AMNH_PBI 00139044) (AMNH). arthropod symbionts (Araneae; Hemiptera;
Diptera) inhabiting webs of the spider Ten-
Acknowledgments gella radiata (Araneae, Tengellidae). Ameri-
can Museum Novitates 3065: 17 pp.
I thank Steve Thurston for assistance
SˇTYSP. (1967): Lipokophila chinaigen.n., sp.n. – a
with inking, scanning, and layout of all il- new genus of Plokiophilidae (Heteroptera)
lustrations in their final digital form. from Brasil. — Acta entomol. Bohemoslov. 64:
Thanks to Jakob Mey, Angela Klauss, Mar- 248-258.
garet Hart, and Christiane Weirauch for as-
Address of the Author:
sistance with scanning electron microscopy.
I especially thank Christiane for assistance Dr. Randall T. SCHUH
with morphological preparations, discussion Division of Invertebrate Zoology
of the relevant literature, interpretation of American Museum of Natural History
morphological structures in Heissophila, and New York, New York 10024
for commenting on a draft of the manu- USA
script. Map prepared by Michael Schwartz. E-Mail: [email protected]
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