See

discussions, stats, and author profiles for this publication at:

https://www.researchgate.net/publication/236155131

Tarsus I chaetotaxy and structure in

Anactinotrichida mites: characters
with phylogenetic value

Article January 2005

CITATIONS READS

4 104

1 author:

Mara Lourdes Moraza

Universidad de Navarra
165 PUBLICATIONS 690 CITATIONS

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Several genere of the family Ascidae View project

Diversity of mites View project

All content following this page was uploaded by Mara Lourdes Moraza on 27 September 2017.

The user has requested enhancement of the downloaded file.

Phytophaga, XIV (2004): 347-359
I S S N: 0393 - 8131

Tarsus I chaetotaxy and structure

in Anactinotrichida mites:
characters with phylogenetic value

MARIA LOURDES MORAZA

Summary

The patterns of structure, chaetotaxy and configuration of the dorso-distal re-

ceptor complex of tarsus I, as found in the larval and subsequent instars of An-
actinotrichida mites allow us to determine homologies among a variety of tarsal
structures and to formulate the following phylogenetic hypothesis: Ixodida is the sis-
ter group of Holothyrida, these two groups in turn form the sister group of the
Mesostigmata. The Opilioacarida is the sister group of the Parasitiformes.

Key words: Acari, Anactinotrichida, tarsus I, structure, chaetotaxy, phylogeny.

Introduction

Tarsus I in Anactinotrichida mites, in addition to the receptor area re-

ferred to as Hallers organ in the Ixodida and Holothyrida, has other tarsal
setae (sensory and solid core setae) whose presence, ontogeny, position, mor-
phology and function may assist in clarifying the phylogeny and classifica-
tion of these mites. While the homology of the Hallers organ has been used
to propose a phylogenetic relationship between Ixodida and Holothyrida
(Lehtinen, 1991), the homology or similarities between this organ and the
receptor complex (CTR) present in Mesostigmata and Opilioacarida re-
mained to be clarified.
The complete chaetotaxy of tarsus I in the larval and postlarval instars,
pore-like structures and setal morphology of Ixodida (Ixodidae and Argasi-
dae) have already been studied by several authors (Clifford and Anastos,
1960; van der Hammen, 1983; Hess and Vlimant, 1986; Leonovich, 1987;
Klompen, 1992; Klompen and Oliver, 1993). The larval instar of Holothyrida
remains undescribed and only the number of verticils of setae on tarsi II-IV
is known (Klompen, 1992); a few papers describe the nymphal and adult in-

Contribution of the 5th EURAAC - Symposium, Berlin 2004.

348 Phytophaga, XIV, 2004

stars (van der Hammen, 1961, 1965, 1968a, 1983, 1989) but none provide a
detailed description of tarsus I chaetotaxy, and illustrations and descriptions
have been confined to the Hallers organ and surrounding areas.
Unpublished observations (Moraza in prep.) of tarsus I in Mesostig-
mata provide a complete pattern of chaetotaxy, ontogeny and morphology
for all groups of Mesostigmatic mites. Other studies have concentrated on
the receptor complex, establishing the homology of several setae with those
in the Hallers organ of Ixodida (Haarlov, 1943; Leonovich, 1984, 1985, 1986,
1987, 1989; Pugh, 1996).
Tarsal chaetotaxy of larval Opilioacarida has been illustrated by Klom-
pen (2002) for Opilioacarus texanus, but their descriptions for adult and
nymphal instars are scarce (van der Hammen, 1966, 1968b, 1977; Juvara-
Bals and Baltac, 1977, etc).
The present paper summarizes the information about structure and
chaetotaxy of tarsus I in Anactinotrichida mites and provides new data with
the hope of improving our understanding of the phylogenetic relationship
between Anactinotrichida mites.
This presentation follows Johnston (1982) in treating the suborder
Opilioacarida in an order, Opilioacariformes, separate from the order Para-
sitiformes. The latter includes the suborders Holothyrida, Ixodida, and
Mesostigmata.

Material and Methods

This work is based on personal observations of described species of Ixo-

dida and Mesostigmata, one undescribed species of Opilioacarida and one
undescribed larva of Holothyrida. Other original descriptions of An-
actinotrichida mites have been carefully analyzed. For setal notation and
terminology, see Grandjean (1942), Evans (1963), Klompen (1992) and van
der Hammen (1976, 1989). In the diagrammatic representations of the
tarsal chaetotaxy, circles represent larval setae; stellate symbol represents
larval phanere with a lanceolated terminal part; rhombuses protonymphal
setae in Mesostigmata; triangles deutonymphal setae in Mesostigmata and
postlarval setae in Opilioacarida, Ixodida and Holothyrida, and solid black
symbols ventral setae. To interpret setal function, the studies of Leonovichs
(1984, 1985, 1987, 1989) and Hess and Vlimants (1986) relating to tarsal
sensilla in different families and genera of Mesostigmata and Ixodida are
taken into account.
Dotted symbols denote absent or missing setae in the illustrated taxa;
solid punctated symbols refer to sw-wp-sensilla, olfactory setae of
Leonovich, (1989) and wp-sw type A and B of Hess and Vlimant (1986); sol-
id diagonal acutely hatched symbols refer to wp-dw/A, solid horizontally
hatched symbols refer to wp-dw/B, solid diagonal back-slashed symbols re-
fer to wp-dw/C, and solid checked symbols to tp-/A,B of Hess and Vlimant
(1986) (see symbols under fig. 6b).
 M.L. Moraza: Tarsus I chaetotaxy and structure in anactinotrichida... 349

The differences in chaetic patterns are shown numerically by the

chaetotactic formula: al d / v pl (number of antero-lateral setae num-
ber of dorsal setae / number of ventral setae number of postero-lateral se-
tae); or al ad/av md/mv pd/pv pl (ad = number of antero-dorsal setae,
av = antero-ventral setae, md = medio-dorsal setae, mv = medio-ventral se-
tae, pd = postero-dorsal setae, pv = postero-ventral setae) (Evans, 1963).
When the notation such as d1 (unitalicized roman notation) are used, they
include a pseudosymmetric pair ad1 and pd1.
The number of verticils is determined depending on the number of
transversal alignments or whorls of setae. When a verticil is complete, the
transversal alignment includes one pair of lateral and at least one pair of
ventral and one pair of dorsal setae (e.g., 1 - 2 / 2 - 1); an incomplete verticil
lacks one or both dorsal, lateral or ventral setae (e.g., 0 - 2 / 0 - 0, 1 - 2 / 1 - 0).
Verticils are simple or complex: simple verticils have a maximum number of
six setae (1 - 2 / 2 - 1) when they are complete; complex verticils have more
than six setae, more than two dorsal and/or more than two ventral setae and
always one pair of lateral setae (e.g., 1 - 4 / 2 - 1, 1 - 2 / 3 - 1, 1 - 5 / 3 - 1).

Results

Tarsus I of Anactinotrichida mites has four clearly defined compo-

nents: ambulacrum, acrotarsus, telotarsus and basitarsus. These compo-
nents exhibit various chaetic and structural patterns.

Acrotarsus I (Figs 2-4, 6-8)

The acrotarsus or secondary segmentation of tarsus I may be present
in Parasitiformes and may have different configurations although sharing
a similar larval dorsal chaetotaxy (figs 2-4, 6-8):
In Allothyridae, considered the most primitive family of Holothyrida
(Lehtinen, 1991), the acrotarsus is present in all instars. In the larva (1 - 9 /
3 - 1), the pit is not present (e.g., fig. 2). In the adult (fig. 6a), a postlarval api-
cotarsus described by van der Hammen (1989) does not include setae asso-
ciated with the anterior pit or not all of them since the seta with a lance-
olated apical part (stellate symbol), which is clearly acrotarsal in the larva,
is found in a groove with other setae. The acrotarsus is always absent in
Holothyridae and Neothyridae (fig. 6b).
The acrotarsal fissure may be present in all instars of several species of
Ixodidae (larva to adult, e.g., A. variegatum), but it is always absent in Ar-
gasidae. When present, some peculiarities should be noted: peripodomeric di-
vision is complete in the larval instar (fig. 3) and it bears dorsal setae of verti-
cil 1 (1 - 7 / 2 - 1); in nymphs and adults, this division may be effaced dorsally
between the pit and the capsule, or the pit may have shifted to a more basal
position (fig. 7) giving rise to the postlarval apicotarsus as in adult Allothyri-
dae which is characterized by the absence of the Hallers organ (1 - 4(6) / 4 - 1).
 350
Phytophaga, XIV, 2004

Figs 1-4 - Diagrammatic representation of tarsus I larval chaetotaxy in Anactinotrichida mites. 1. Larva of Opilioacarus texanus based on Klompen
(2000); 2. Larva of undescribed species of Allothyridae, setal notation based on van der Hammen (1989); 3. Larva of Ixodidae, setal notation modified
from Klompen (1992); 4. Larva of Mesostigmata (Sejus), setal notation modified from Evans (1963).
 M.L. Moraza: Tarsus I chaetotaxy and structure in anactinotrichida... 351

The acrotarsus is present in all instars of Sejina (considered the earli-

est derivative group of mesostigmatic mites by Johnston (1982), Evans
(1992), Athias-Henriot, (1972), Lehtinen (1991)) and in Heterozerconina. In
the larva (1 - 7 / 3 - 1), the pit is not present (e.g., fig. 4) and there are three
ventral setae as in the larva of Allothyridae. The postlarval acrotarsus of
these mites is similar to the larval one, except for the postlarval setae that
complete the acrotarsal chaetotaxy (fig. 8). In Parasitina and Dermanyssi-
na, the acrotarsus may appear in the deutonymph and this derivative acro-
tarsus includes some dorsal telotarsal setae.
In Opilioacarida, the acrotarsal articulation is effaced in all instars,
such as in tarsi II-IV in the larval instar.

Telotarsus I
Telotarsus I (including the acrotarsus) in Parasitiformes exhibits two
main larval conditions:
* In the larva of Allothyridae tarsus I has nine complex verticils of setae
(8 - 51 / 15 - 8) (fig. 2): V1 (1 - 9 / 1 -1), V2 (1 - 15 / 2 - 1), V3 (1 - 3 / 2 - 1), V4-V6
and V8 (1 - 4 / 2 - 1), V7 (1 - 5 / 2 - 1) and V9 incomplete (0 - 3 / 0 - 0). The three
distal verticils bear hollow sensorial setae (receptor complex CTR); the six
basal verticils bear solid tactile setae (one solenidion may be present). In
adults and nymphs (fig. 6a), at least 13 tarsal verticils are present and each
verticil bears four or five dorsal setae and either two or four ventral setae.
* In the larva of Ixodida and Mesostigmata (figs 3, 4), there are four, al-
beit incomplete, ancestral verticils. Verticils 1 and 2 are complex and have,
together with verticil 3, some hollow sensorial setae (CTR), while basal ver-
ticil 4 is chaetal (typically solid, with socket, and usually ornate tactile se-
tae) and simple. Both groups bear a maximum of 19-20 dorsal larval setae.
In the larva of Ixodidae (fig. 3), verticils are arranged as follows: V1 (1 -
7 / 2 - 1) with setal types np/C,D,E and wp (porose olfactory setae), V2 (1- 1/1
6/0 1 / 1 - 1) with wp setae, V3 (1 - 2 / 2 - 1) with setae type tp/A (gustatory)
and V4 (0 - 2 / 0 - 0) with setae type np/B (functional terminology of Hess and
Vlimant, 1986). These four larval telotarsal verticils may replicate in the
postlarval instars. In adults of A. variegatum (fig. 7) the nine or teen verticils
are as follows: V1 (1 - 10 / 2 - 1) adding gustatory tp/A setae, V2 (1 - 1/1 11/0
1/1 - 1) with more olfactory setae wp, V3 x 3 (3 - 3/3 2/3 - 3) adding gustatory
tp/A setae and V4 x 4 (4 - 4/4 4/4 - 5). Larval verticils 1 and 2 do not replicate
lateral setae; larval verticil 2 does not replicate ventral setae. Ixodidae al-
ways retain the replication of lateral and ventral setae on verticils 3 and 4,
although dorsal chaetotaxy may remain larval (e.g., B. microplus). Adult Ar-
gasidae ticks usually retain the ancestral telotarsal complement of larval se-
tae and some species show larval supresion of lateral proximal setae.
In the larva of Mesostigmata (fig. 4), the verticil arrangement is simi-
lar to Ixodidae except for V1 (1 - 1/1 5(6)/0 0/0 - 1) with only one ventral se-
ta (ventral setae of V2 are included in the acrotarsus) and with olfactory se-
tae wp and gustatory seta tp; V2 (1- 1/1 5(6)/0 1/1 -1) with olfactory setae wp
 352
Phytophaga, XIV, 2004

Figs 5-8 - Diagrammatic representation of tarsus I chaetotaxy in adult Anactinotrichida mites. 5, Opilioacarus sp., setal notation follows Juvara and
Baltac (1977); 6a. Holothyrida, Australothyrus ocellatus based on van der Hammen (1989); 6b. Holothyrida, Holothyrus coccinella, distal region of tar-
sus I based on van der Hammen (1989); 7. Ixodida, Amblyomma variegatum based on Hess and Vlimant (1986); 8. Mesostigmata: 8a, Sejus, 8b. Dia-
grammatic representation of distal verticils V1 and V2 of Trigynaspida (Cercomegistus).
 M.L. Moraza: Tarsus I chaetotaxy and structure in anactinotrichida... 353

(sw and dw), and V4 complete (1 - 2 / 2 - 1) with solid setae. The complement
of larval setae (4 - 17 / 7 - 4 in Sejina and Heterozerconina and 4 - 19 / 7 - 4
in other Mesostigmata) adds 12 setae in the protonymph and eight setae in
the deutonymph (e.g., fig. 8) (5 - 26 / 8 - 5 in Sejina and Heterozerconina and
5 - 35 / 9 - 5 in other cohorts). Adult verticils are arranged as: V1 (1 - 1/1
9(12)/0 1/1 - 1), V2 (1 - 1/1 6(9)/0 1/1 - 1), V3 (1 - 1/1 4/0 1/1 - 1) with hollow
setae of unknown function and solid verticils V4 (1 - 2 / 0(1) - 1) and V5 (1 -
2 / 2 - 1) (fig. 8a). These mites do not exhibit replication of larval lateral and
ventral setae in verticils 1 to 3 (only replicate dorsal setae and complete the
ventral pair of V1) and only larval verticil 4 duplicates lateral, ventral and
dorsal setae; only one deutonymphal ventral seta (mv) in medial position
augments the four larval pairs of ventral setae.
In the larval condition of Opilioacarida (Klompen, 2000) (fig. 1) the pat-
tern of distribution of setae is not so clear. It appears that the same five
telotarsal verticils of tarsi II-IV are present on tarsus I, with a dorsal com-
plex chaetotaxy. The larval arrangement may be: V1 (1 - 9 / 2? - 1), V2 (1 - 11
/ 3? - 1), V3 (0 - 5 / 2? - 1), V4 (0 - 4 / 2 - 1) and V5 (0 - 3 / 2 - 1). In adults (e.g.,
Opilioacarus sp., fig. 5), the number of telotarsal+acrotarsal verticils is
eight and these verticils are complex: V1 (1 - 11 / 5 - 1), V2 (1 - 3/1 9/1 3+2/1
- 1), V3 (1 - 2/1 3/1 1/1 - 1), V4 (1 - 2/1 4/1 2/1 - 1), V5 (1- 2/1 4/1 2/1 - 1), V6 (1-
1/1 2/1 1/1 - 1), V7 (1 - 2/1 1/2 2/1 - 1) and V8 (1 - 1/1 0/1 1/1 - 1). Hollow sen-
sorial setae are located on the two most apical verticils and in the next ver-
ticils distal to the ornamentation line which separates the smooth tarsal
distal surface from the ornate basal surface.

Basitarsus I
In opilioacarids (fig. 1), the adesmatic basal division is complete in all
instars and the basitarsus and telotarsus of leg I are separated ventrally by
two lyrifissures, laterally and dorsally by soft cuticle (van der Hammen,
1966). In the larva, the basitarsus has four verticils of setae (4 - 4/4 5/3 - 4)
(Klompen, 2000) and the adult exhibits a similar neotrichy to that on ba-
sitarsi II-IV and has one dorsal lyrifissure (e.g., 7 - 6/3 2/3 7/3 -7, fig. 5).
Among Parasitiformes, the adesmatic division is generally incomplete,
allowing very little or no mobility. It is sometimes absent altogether. They
exhibit various conditions:
* In Holothyrida, the basitarsal peripodomeric fissure is always absent
(fig. 2). However, basitarsal hypertrichy is evident, and includes a conjec-
tural number of verticils of dorsal, lateral and ventral setae.
* In Ixodida (fig. 3), only the ventral fissure with lyrifissure itav is pre-
sent and the basitarsus always lacks setae (0-0/0-0).
* In Mesostigmata (fig. 4), a complete or incomplete peripodomeric fis-
sure formed by the entire or partial coalescence of ventral and dorsal lyri-
fissures is present in the proximal region of the podomere. Basitarsus I
lacks setae (0 - 0/0 - 0) in all instars (except for replicate pair l4 in the deu-
tonymph and in the adult Cercoleipus 1- 0/0 - 1).
354 Phytophaga, XIV, 2004

Receptor Complex
A diversified receptor complex is found in all groups of Anactinotrichi-
da mites. However, only in Ixodida and Holothyrida are some of the sensor-
ial setae of this complex associated with a well-differentiated and homolo-
gized Hallers organ.
Opilioacarida mites (figs 1, 5) exhibit the following receptor complex:
an acrotarsal sensorial region with a maximum number of 11 sensorial dor-
sal setae (verticil 1); a cluster of eight olfactory setae, seven of them post-
larval (four may be located in a limited circular area in Opiliacarus vander-
hammeni, Juvara-Bals and Baltac, 1977) at the distal level of larval and
postlarval verticil 2; a posterolateral telotarsal organ (small postlarval cap-
sule with two larval setae) at the basal level of postlarval verticil 2. Other
olfactory setae (two larval ones denoted as T6) are inserted at basal level
of verticil 2 in larva and at basal level of verticil 3 in adult and dorsal sen-
sorial setae of the next basal verticils are on the smooth telotarsal surface
(in Opilioacarus texanus and Paracarus hexophthalmus, the number of ol-
factory setae in the cluster of V2 is reduced and other olfactory setae are lo-
cated in the next basal telotarsal verticils, postlarval verticil 4 and others).
This type of arrangement is considered to be the primitive condition of the
receptor complex in Anactinotrichida mites (van der Hammen, 1983; Lehti-
nen, 1991) as follows: acrotarsal region without a pit; telotarsal distal region
without a capsule containing a cluster of olfactory setae; telotarsal medial
region with a small postlarval capsule in posterolateral position and with
smooth surface with several porose setae. The basal telotarsal region is or-
nate and bears only solid setae.
Adult Holothyrida and Ixodida (figs 6, 7) show the most derivative and
specialized condition: a group of acrotarsal setae in a groove or pit or de-
pression (anterior pit) and the distal telotarsal region with another group of
setae clustered in a new derivative capsule.
In larval Allothyridae (fig. 2), the pits setae are on the surface of the
acrotarsus and the capsules setae are on the distal region of the telotarsus
although the pit and capsule themselves are absent. The two setae denoted
as 5 and 6 retain the primitive basal position on V2 together with two hol-
low setae (d0) which delimit the basal telotarsal region. In adult Allothyri-
dae (fig. 6a), the dorsal groove is in a mediodistal telotarsal position (such as
in Ixodida with apicotarsus) and includes three setae, and setae 5 and 6
are included in the postlarval posterolateral capsule together with seven
other setae (based on the illustration provided by van der Hammen
(1989)). In Holothyridae (fig. 6b) and Neothyridae, which both lack an acro-
tarsus in all instars, the same nine capsular setae are found in a circular
medial depression that resembles the circular cluster in Opilioacarus van-
derhammeni and passes anteriorly into an elongated groove ending be-
tween the two terminal apophyses; other sensorial acrotarsal setae are po-
sitioned alongside this groove (the groove is absent in the smallest known
nymphal instar of Holothyridae (van der Hammen, 1983)).
 M.L. Moraza: Tarsus I chaetotaxy and structure in anactinotrichida... 355

In the larva of Ixodida (figs 3), an acrotarsal pit with five setae and a
telotarsal distal capsule with four or five setae are present; two setae de-
noted as dm2 of the telotarsal region (wp-dw/A such as in most Mesostig-
mata) remain in the ancestral basal position and may be located in a small
medial depression in Argasidae, and two setae d3 with porose tip (tp/A)
are in V3. In adult Ixodida (fig. 7), one seta may be added in the pit, three or
in the capsule, two more setae dm2 in the telotarsal basal region, and the
sensorial setae of V3 may replicate. In some species, the acrotarsal pit and
capsule may share a common depression (e.g., Ixodes ricinus).
Larval and adult mesostigmatic mites exhibit a similar condition (figs
4, 8) as larval Allothyridae (fig. 2): acrotarsal pit and telotarsal capsule ab-
sent; lanceolated phanere and four or five larval and five to seven postlarval
porose setae on the dorsal acrotarsal surface; disto-medial region of verticil
2 with olfactory setae in a cluster (T1 and T2 may be in a small capsular de-
pression) surrounding the acrotarsal sensorial setae (e.g., Davacarus, Cer-
comegistina) or they may be more or less dispersed on the tarsal surface and
basal to verticil 2; telotarsal basal region of verticil 2 with two (T6), three
or four (T7, T8) porose setae basal to or at the same level as setae d2 and
hollow setae of V3 (two larval and four postlarval).

Homologies in the Receptor complex

Based on their ontogeny, their initial (larval) and final (adult) topo-
graphic position on the surface of the segment (slight changes of the topo-
graphic position through ontogeny usually occur among most setae), their
morphology and function (and considering that morphology and function
may change through ontogeny in several cases), the following setal/phanere
homologies are proposed. In Mesostigmata, apical setae ad1 and md1 ap-
pear homologous to both setae dm1 in Ixodidae; ad1 (and non-porose dm1 in
Ixodidae) to seta dy in Holothyrus; seta J (porose seta, phanere with a
lanceolated terminal part also found in Allothyridae) may be denoted z as
in Holothyrus and as c (conical setae) in Ixodida; setae J2 and J3 may be ho-
mologous to 1 and 2 in Holothyridae and to setae f in Ixodida; setae A1
may be homologous to one p (porose wp-sw/A) and A2 may be homologous to
g (porose grooved setae) in Ixodida and to two capsular porose in
Holothyrida. Five distal telotarsal setae T1-T5 may be homologised with
five of the capsular setae in Ixodida and Holothyrida. Two more postlarval
acrotarsal setae in Mesostigmata A6, A7 may be homologous to two of
the seven or nine capsular setae in Ixodida and Holothyrida, suggesting
that the dorsal proximal limit of the acrotarsus may change (such as in the
deutonymphal acrotarsus), and as capsular setae are not confined within
the capsule, their proximal or distal position in relation to the acrotarsal fis-
sure may change as well. Basal telotarsal setae T6 may be homologous to
denoted 5 and 6 in the larva of Allothyridae and to dm2 in Ixodida.
The acrotarsal origin hypothesised by van der Hammen (1961) for the
setae associated with the anterior pit appears confirmed, since they are al-
356 Phytophaga, XIV, 2004

so found on the larval acrotarsus of all Parasitiformes mites even when the
pit is absent. Regarding the telotarsal sensory area, it appears that the cap-
sular structure, larval in Ixodida but postlarval in Allothyridae as in Opil-
ioacaridae, is of telotarsal origin. When the capsule is not present in the lar-
va the setae included therein may have a dual origin, acrotarsal and
telotarsal (such as in Allothyridae). In Opilioacarida, the postlarval capsule
includes only two telotarsal setae of the central telotarsal region of V2.

Discussion and Conclusions

The recorded variability in structure, chaetotaxy and other character-

istics of tarsus I among the main taxa of Anactinotrichida mites allow us to
propose significant hypotheses regarding homologies of shared structures
and phylogenetic relationships among these mites.
The receptor complex has four ancestral regions: acrotarsal (verticil 1),
telotarsal distal region of larval verticil 2, telotarsal medial region of larval ver-
ticil 2 and telotarsal basal region of verticil 2. The first two regions with the de-
rivative acrotarsal pit and the capsule have been defined as Hallers organ.
The presence of nine larval verticils of setae on tarsus I of allothyrids,
much as in Opilioacarida, with the complete absence of a basitarsal ring,
and the presence of five or six larval verticils of setae on tarsi II-IV (Klom-
pen, 1992), two of them on the basitarsi (Klompen, pers. comm.), lead us to
believe that at least two larval verticils belong to the ancestral basitarsus I
although with a newly derivative replicate condition.
Notable symplesiomorphies link Holothyrida with Opilioacarida: the
number of tarsal verticils of setae in the larva, retention of at least five an-
cestral telotarsal verticils, the posterolateral position and postlarval on-
togeny of the telotarsal capsule in Allothyridae and the circular cluster of ol-
factory disto-telotarsal setae in Holothyridae when the capsule is open.
Tarsus I in larval and adult holothyrids and opilioacarids has complex ver-
ticils, with more than six setae. However, the ancestral larval condition on
legs I-IV is different in the two groups: the larva of holothyrids has complex
verticils on individed tarsi I-IV and this apparently derived condition is re-
tained in postlarval instars; opilioacarids exhibit larval and postlarval sim-
ple verticils on movable basitarsi I-IV and on telotarsi II-III (the latter con-
dition shared with Ixodida and Mesostigmata).
Ixodida is regarded as the sister group of Holothyrida based on the
shared derivative condition of the receptor complex (an acrotarsal anterior
pit and a telotarsal capsule including the telotarsal distal olfactory setae
(C1-n) in Ixodida and including the distal (1-4) and probably basal
telotarsal sensorial setae (5 and 6) in Holothyrida), absence of dorsal ba-
sitarsal fissure and lyrifissure itd and derivative postlarval acrotarsus (api-
cotarsus). As plesiomorphies, Holothyrida and Ixodida share a replication of
tarsal verticils. Ixodida exhibits the derivative condition of possessing a lar-
 M.L. Moraza: Tarsus I chaetotaxy and structure in anactinotrichida... 357

val pit and capsule; the ontogenetic retardation of setae v4 and basitarsal
setae l4 on tarsus I is shared with Allothyridae. Complex larval verticils on
basitarsal region of tarsus I, the lack of a basitarsal fissure and basitarsal
ventral lyrifissure on tarsus I are synapomorphic characters for the
Holothyrida. The postlarval posterolateral capsule of Allothyridae is proba-
bly independently derived between Holothyrida and Opilioacarida.
Holothyrida + Ixodida is regarded as the sister group of Mesostigmata,
in support of the position held by Lehtinen (1991) and opposing the scheme
of relationships presented by Norton, Kethley, Johnston and OConnor
(1993); the latter assumed, without a discussed basis, that Holothyrida is
the sister group of Mesostigmata, and that Holothyrida + Mesostigmata is
the sister group of Ixodida. Mesostigmata retain the ancestral larval condi-
tion of the receptor complex of Allothyridae, including the ancestral mid-lev-
el position of setae T6 of Opilioacarida, the simple verticils on basitarsal
region of tarsus I of Ixodida and Opilioacarida and probably the dorsal ba-
sitarsal lyrifissure itd present on Opilioacarida. The reduced nude basitar-
sus I shared with Ixodida may be an apomorphic condition perhaps inde-
pendently derived. The presence of the deutonymphal acrotarsus is a
derivative characteristic of Mesostigmata mites.
In comparison with Opilioacarida, the reduction or complete lack of
adesmatic basitarsal division on tarsus I, a receptor complex confined to the
three distal tarsal verticils, and the presence of an acrotarsus I in Parasiti-
formes are derivative characters (synapomorphies) which distinguish these
mites from the Opilioacarida, the latter with their long movable neotrichous
basitarsus I and lack of an acrotarsus I. Synapomorphic attributes of tarsi II
to IV also distinguish Parasitiformes from Opilioacarida (Lindquist, 1984).
The numerous cuticular receptors clustered on the dorsoapical surface of tar-
sus I is a character complex shared among Opilioacarida, Holothyrida, Ixo-
dida and Mesostigmata, and this attribute may be one of the basic apomor-
phies characteristic of the Anactinotrichida (Lindquist, 1984).

Acknowledgements

Special thanks to Dr. Evert Lindquist for his generosity and confidence in pro-
viding me with the Opilioacarida materials and his discussions and review of drafts
of this paper, and to my teacher, colleague and dear friend Dr. Donald Johnston, who
provided the inspiration for this work.

References

ATHIAS-HENRIOT C. - 1972 - Gamasides chiliens (Arachnides). II. Rvision de la famille

Ichthyostomatogasteridae Sellnick, 1953 (=Uropodellidae Camin, 1955). - Archos
Zool. Est. S Paulo, 22: 113-191.
CLIFFORD M.C., ANASTOS G. - 1960 - The use of chaetotaxy in the identification of larval
ticks (Acarina: Ixodidae). - Jour. Parasitol. 46 (5): 567-578.
358 Phytophaga, XIV, 2004

EVANS G.O. - 1963 - Observations on the chaetotaxy of the legs in the free-living Gamasi-
na (Acari: Mesostigmata). - Zool. Bull. Br. Mus. Nat. Hist., 10: 277-303.
EVANS G.O. - 1992. Principles of Acarology. Wallingford: C.A.B. International, 563 pp.
GRANDJEAN F. - 1942 - Observations sur les Acariens (7e srie). - Bull. Mus. Nat. Hist.
Nat., 2 (14): 216-220.
HAARLOV N. - 1943 - The chaetotaxis of Tarsus I in some mesostigmata. With special
reference to a supposed homologon of the Tarsus organ of the genus Spinturnix
and Hallers organ of the ticks. - Entomol. Meddelelser, 23: 273-294.
HESS E., VLIMANT M. - 1986 - Leg sense organs of ticks. - In: J.R. Sauer, A. Hair (eds)
Morphology, physiology and behavioral biology of ticks, Ellis Horwood, Chichester.
361-390.
JOHNSTON D.E. - 1982 - Parasitiformes. - In: S.B. Parker (ed) Synopsis and Classification
of Living Organisms, McGraw-Hill, New York, vol. 2: 111-117.
JUVARA-BALS I., BALTAC M. - 1977 - Deux nouvelles espces dOpilioacarus (Acarina:
Opilioacarida) de Cuba. In: T. Orghidan, A. Nnez Kimnez A, V. Decou, ST. Ne-
grea, N. Via (eds) Rsultats des expditions biospologiques cunano-roumaines
Cuba, 172
KLOMPEN J.S.H. - 1992 - Comparative Morphology of Argasid Larvae (Acari: Ixodida: Ar-
gasidae), with Notes on Phylogenetic Relationships. - Ann. Entomol. Soc. Amer., 85
(5): 541-560.
KLOMPEN J.S.H. - 2000 - Prelarva and larva of Opilioacarus (Neocarus) texanus (Cham-
berlin and Mulaik) (Acari : Opilioacarida) with notes on the patterns of setae and
lyrifissures. - Journal of Natural History, 34 (10): 1977-1992.
KLOMPEN J.S.H., OLIVER J.H. - 1993. Hallers organ in the tick family Argasidae (Acari:
Parasitiformes: Ixodida). - J. Parasitol., 79 (4): 591-603.
LEHTINEN P.J. -1991 - Phylogeny and zoogeography of the Holothyrida. - In: F. Dusbabek,
V. Bukva (eds) Modern acarology, SPB Academic Publishers, The Hague, Vol. 2:
101-113.
LEONOVICH S.A. - 1984 - Tarsal receptor complexes of gamasid mites of the family
Haemogamasidae. - Parazitologiya, 18 (6): 451-453.
LEONOVICH S.A. - 1985 - Ultrastuctural analysis of the tarsal receptor complex of gamasid
mite Hirstionyssus criceti (Hirstionysisdae). - Parazitologiya, 19 (6): 456-463.
LEONOVICH S.A. - 1986 - Tarsal receptor complexes of gamasid mites of the family
Haemogamasidae. - Parazitologiya, 18 (6): 451-453.
LEONOVICH S.A. - 1987 - The search receptors of bloodsucking ticks and mites (Parasiti-
formes). - Parasitol. Sbornik, 34: 83-79.
L EONOVICH S.A. - 1989 - Tarsal receptor complex and systematics of gamasid mites
(Parasitiformes, Mesostigmata, Gamasina). - Parazitologiya, 23 (6): 469-479.
LINDQUIST E.E. - 1984 - Current theories on the evolution of major groups of Acari and on
their relationships with other groups of arachnida, with consequent implications for
their classification. - Acarology, 6 (1): 2862.
NORTON R.A., KETHLEY J.B., JOHNSTON D.E., OCONNOR B.M. - 1993 - Phylogenetic per-
spectives on genetic systems and reproductive modes of mites. - In: D.L. Wrensch,
M.A. Ebbert (eds) Evolution and diversity of sex ratio in insects and mites, Chap-
man and Hall, New York. pp. 8-99.
PUGH P.J.A. - 1996 - The structure and function of the Tarsus I sensillar field in mites of
the genus Halarachne (Halarachnidae: Gamasina). - J. Nat. Hist., 30: 1069-1086.
VAN DER HAMMEN L. - 1961 - Description of Holothyrus grandjeani nov. spec., and notes on
the classification of the mites. - Nova Guinea, Zool., 9: 173-194
VAN DER H AMMEN L. - 1965 - Further notes on the Holothyrina (Acarina) I. Supple-
mentary description of Holothyrus coccinella Gervais. - Nova Guinea, Zool., 40
(28): 253-276.
VAN DER H AMMEN L. - 1966 - Studies on Opilioacarida (Arachnida) I. Description of
Opilioacarus texanus (Chamberlin & Mulaik) and revised classification of the
genera. - Zoologische Verhandelingen, Leiden, 86: 1-80.
 M.L. Moraza: Tarsus I chaetotaxy and structure in anactinotrichida... 359

VAN DERH AMMEN L. - 1968a - Stray notes on Acarida (Arachnida) I. - Zoologische

Mededelingen, Leiden, 42 (25): 261-280.
VAN DER HAMMEN L. - 1968b - Studies on Opilioacarida (Arachnida) II. Redescription of
Paracarus hexophthalmus (Redikorzev). - Zoologische Mededelingen, Leiden, 43 (5):
57-76.
VAN DER HAMMEN L. - 1969 - Studies on Opilioacarida (Arachnida) III. Opilioacarus
platensis Silvestri, and Adenacarus arabicus (With). - Zoologische Mededelingen,
Leiden, 44 (8): 113-131.
VAN DER HAMMEN L. - 1976 - Glossaire of acarological terminology. Vol. II Opilioacarida. -
Dr. W. Junk Publishers. 137 pp.
VAN DER HAMMEN L. - 1977 - Studies on Opilioacarida (Arachnida) IV. The genera Pan-
chaetes Naudo and Salfacarus gen. nov. - Zoologische Mededelingen, Leiden, 51 (4):
43-78.
VAN DER HAMMEN L. - 1983 - New notes on Holothyrida (Anactinotrichid mites). - Zoolo-
gische Verhandelingen, Leiden, 207: 1-48.
VAN DER HAMMEN L. - 1989 - Glossaire of acarological terminology. Vol. III Holothyrida. -
Accademic Publishing bv, The Hague, 116 pp.

Authors address
M.. MORAZA - Departamento de Zoologa y Ecologa, Facultad de Ciencias, Universidad
de Navarra, C/ Irunlarrea s/n, Pamplona 31080, (Navarra) (Spain).
View publication stats