Phylogeny of the genera of Ticoplinae (Hymenoptera: Mutillidae)

14 Pages·2002·6.9 MB·English

by  Andrew Mitchell

HYM. RES.
J.
Vol. 11(2), 2002, pp. 312-325
Phylogeny of the Genera of Ticoplinae (Hymenoptera: Mutillidae)
Andrew Mitchell and Denis Brothers
J.
Sche^ol of Botany and Zoology, University of Natal Pietermaritzburg, Private Bag XOl,
Scottsville, 3209 South Africa (AM now: School of Molecular and Cellular Biosciences);
AM
email: [email protected] DJB email: [email protected]
—
Abstract. The subfamily Ticoplinae Nagy is one of the more basal taxa in Mutillidae. Cladistic
analyses using 21 characters have demonstrated that recognition c^f two tribes is supported. Ti-
coplini Nagy includes Nmiomutilla Andre (= Ticopla Nagy syn. nov.) and Arcotilla Bischoff; Smi-
cromyrmillini Argaman includes SDiicroiinfiiiiilla Suarez (and possibly Cniiicroiiilla Lelej and Krom-
bein, Eosiiiicwmi/nfiilla Lelej and Krombein, and Hiudiistaiulln Lelej and Krombein, should these
be considered valid). The subfamily and tribes are reviewed and characterized, a key to tribes
and genera is provided, and bothsexesoftypical membersofthethree maingeneraareillustrated.
Nanoinutilla nadae Argaman 1988 is selected as the correct spelling for the species also spelled N.
nada in its original description, and Areotilla ferniginea Mitchell and Brothers 1998 for the species
also spelled Arcotilla fcrritgi)iata in its original description.
Ticoplinae Nagy 1970 is one of the rel- that they belong in a single subfamily, the
atively basal subfamilies of Mutillidae, as valid name of which is Ticoplinae (and
shown by Brothers (1975, 1999) (Fig. 1) would remain so even if Ticopla were con-
and by Lelej and Nemkov (1997) whose sidered a junior synonym of Nanomntilla;
analyses differed from Brothers' in several International Cc^nimission on Zoological
respects. It was established as a subfamily Nomenclature 1999: Article 40.1). He also
of Nagy's Heterogynidae (properly Het- conclucied that the relationships amongst
erogynaidae; International Commission the component genera were such as to
on Zoological Nomenclature 1987), to ac- preclude the recognition of tribal divi-
commodate the genus Ticopla Nagy 1970, sions. However, Argaman (1988) pro-
known only from male specimens. Broth- posed such divisions: Ticoplini, including
ers (1975) transferred Ticoplinae to Mutil- Ticopla and Nanoinntilla, which he consid-
lidae, and Day (1984) placed Heterogi/iui ered distinct, and Smicromyrmillini, in-
Nagy in Sphecidae s.l.; it is now consid- cluding Siiiicroiin/nnilla. He did not ex-
ered to comprise a distinct family, Heter- amine Arcotilla.
ogynaidae (Brothers 1999, Melo 1999). The The cladistic study presented here elu-
genera placed in Ticoplinae by Brothers cidates the phylogeny of the genera of Ti-
(1975) were Arcotilla Bischoff 1920, Nano- coplinae, enabling objective assessment of
inutilla Andre 1900, Sinicroim/nnilla Suarez Argaman's tribal divisions. The morpho-
1965 and Ticopla. Independently, Suarez logical terms used are those of Gauld and
(1975) proposed a new subfamily, Nano- Bolton (1988). Specimens examined are in
mutillinae, to contain Nanonnitilla, and the collection of one of the authors (DJB)
placed Sniicroiin/nnilla in Myrmillinae but or were borrowed from numerous insti-
he did not realise that Ticopla or Arcotilla tutions (particularly the Natural History
were of relevance. Brothers' (1975) study Museum (London) and Museum National
settled much of the controversy over the d'Histoire Naturelle (Paris)) over many
classification of these genera by showing years.
Volume 11, Number 2, 2002 313
Myrmosinae with their similarity in size, convinced
him that the two specimens were conspe-
Pseudophotopsidinae
cific, despite the fact that they were not
Ticoplinae captured in copula. We have examined the
type series of M. (N.) microsoma in the
Rhopalomutillinae
Transvaal Museum and found that the fe-
Sphaeropthalminae male specimen was actually collected at
Bothaville, in the Free State, while the
Mutillinae male was collected near Willowmore, in
Myrmillinae the Eastern Cape, approximately 700 km
away. They are not conspecific, nor even
Fig. 1. Phylogeny ofsubfamilies of Mutillidae (sim- congeneric; Nonveiller (1973) concluded
plified from Brothers 1975, 1999).
that the male really belongs in Sinicrouiyr-
milla. To add to the confusion, Bischoff
GENERA OF TICOPLINAE (1921) described six new species of Naiio-
niutilla without examining the type spe-
Nauomutilla Andre
cies; all of his new species would later be
(Figs. 2-9)
recognised as belonging in Smicromyrmilln
Mutilhi {Nauoimitilla) Andre 1900: 130. (Nonveiller 1973). Not surprisingly, Ar-
Type species: Miitilla vniidieri Tournier 1895, nold (1946, 1960) made the same error in
Morocco, by subsequent designation of Ash- describing another two species. Nonveiller
mead (1903). (1973) transferred to Sinicrounjrmilln all
Ticopla Nagy 1970: 85.
species ofNauomutilla, except forN. vauch-
Type species: Ticopla yoca Nagy 1970, Jordan,
cri and the female ofN. microsoma, and de-
by original designation. Syn. nov.
limited both genera, although he had also
The first description of Nmioniiitilla ap- not examined the type specimen(s) of N.
peared in a key to subgenera of Mutilln microsoma. We confirm Nonveiller's con-
published in April 1900, withoutany nien- clusions as correct. Argaman (1988) de-
tion of included species. Later, in the same scribed a third species, N. iiadac from
work, Andre (1901b: 223) presented a for- Spain, again known only from female
mal description based on a single species specimens. (Although the name is mostly
known from females only {Mufilhi vniicli- spelled "nada" in that paper, it is "uadac"
cri) but also provided a description of the in the key; there is a statement that the
male in a footnote (p. 224), based on a sec- species is named after Mrs Nonveiller, us-
ond species supposedly known from both ing her nickname [which is Nada, DJB
sexes {Mutilln uiicrosouin) which he had re- pers. obs.], so the feminine genitive form
cently described; it must thus be conclud- is preferable, and the commoner spelling
ed that there were two originally-included is probably an inadvertent error.)
species, although subsequent authors Many statements by other authors re-
have considered NanoDiutilla to have been ferring to 'Nauomutilla haxe been based on
a monotypic genus at establishment. An- a presumption that they apply to the type
dre (1901a) ciescribed both sexes ofMutilln species, N. vaucheri. It is now clear to us,
{Naiioiiiutilhi) niicrosoina from South Africa, however, that the specimens identified
being under the impression that the male and illustrated as N. vaucheri by Non\eill-
and female specimens had been collected er (1973), Argaman (1988), and possibly
in the same area, Willowmore (". . . j'ai Suarez (1975), were misidentified, al-
trouve une autre espece du meme sous- though they were also collected in Moroc-
genre . . . accompagnee d'un male rencon- ct^ (the type locality of N. vaucheri being
tre dans les memes parages"). This, along Tangier). When compared with the origi-
314 Journalof Hymenoptera Research
pir.MiiwH
Figs. 2-9. Naiuvuutilla spp., dorsal and lateral views. 2-5, N. •oniichcii (Tournier), V, length = 2.0 mm ((.ii-
braltar, compared with ht)lotype). 6-Q, N. sp., 6, length = 4.4 mm (Zimbabwe). Scales = 1.0 mm.
nal description (Tournier 1895) and the and Argaman (1988) show the mesosoma
fuller description and illustration by An- as much less slender, without a complete
dre (1901b), some discrepancies are obvi- longitudinal carina and without a median
ous. Both Tournier and Andre referred to posterior tooth; Suarez (1973) expressed
a median longitudinal carina on the me- puzzlement at the lack of such a carina in
sosoma (this is shown in Andre's illustra- specimens he identified as N. vniiclwri but
tion as ending in a fine tooth posteriorly) provided no illustrations. One of us (DJB)
and also stated (and illustrated) that the has examined the holotype of N. vauchcri
mesosoma was twice as long as wide. The (collected at Tangier by Vaucher, with
illustrations given by Nonveiller (1973) Tournier's determination label referring to
Volume 11, Number 2, 2002 315
the publication of the name, labelled as others, one from Algeria and two from Sy-
from the Tournier Collection and housed ria, represent two different species.)
in the Geneva Museum). Unfortunately, it Ticoplii was described for two new spe-
has been glued dorsal-side down to a card cies collected in the Jordan region and
so that the dorsal surface of the mesosoma known only from males (Nagy 1970).
is almost entirely obscured. There is, how- Brothers (1975) suggested thatone ofthese
ever, a clearly conspecific specimen in the may be the male of N. vaucJieri, suppos-
same collection, also collected at Tangier edly known from the same area but only
(in 1896), which is essentially identical to from females (based on specimens so
the holotype (although with the tibiae identified by Andre, see above). Argaman
very slightly paler); the mesosomal dorsal (= Nagy) (1988) countered this by describ-
surface is clearly visible and shows an al- ing females of both species of Ticopla.
most complete very fine median longitu- Nonetheless, he stated that "the resem-
dinal carina ending in a very small pos- blance between Nniioinutilln and Ticopla fe-
terior tubercle, and the mesosoma is rela- males is so remarkable, and the difference
tively more elongate than in the speci- so delicate, [that it is] entirely understanci-
mens illustrated by Nonveiller (1973) and able" that Andre had identified a speci-
Argaman (1988). Another specimen, from men from Syria (that Argaman called a Ti-
Gibraltar and housed in the Natural His- copla) as N. vaiichcri; i.e., Argaman (1988)
tory Museum, London (illustrated here. stated that these genera are so similar as
Figs. 2-3), is also clearly conspecific al- to be easily confused.
though the appendages are slightly paler We have examined five specimens
than in the holotype; it has lost the scat- (three females from Syria and two males
tered long erect setae on the mesosoma from Amman, Jordan) that are unques-
and most of the decumbent pubescence, tionably Ticopla based on Argaman's
but shows the carina and tubercle more (1988) criteria, and find no consistent dif-
clearly as a result. The carina is extremely ferences between them and female speci-
fine and somewhat irregular, normally mens of 6 further species from Gibraltar
concealed under fairly dense diagonally and Morocco {N. vaiichcri) and southern
oriented decumbent pubescence that gives Africa (including N. inicrosonia), and males
the appearance of a mid-dorsal line in un- of 17 species from Kenya and Angola to
worn specimens, and even when visible southern Africa. The differences in the
needs careful illumination; Suarez (1975) sculpturing of the niesosomal dorsum of
may thus have overlooked it, although he both sexes, the main character used by Ar-
did comment on the pubescent line. Both gaman in distinguishing these two genera,
species of Naiioiiiutilln illustrated and dis- are by no means as distinct as he sup-
cussed by Argaman (1988) are different posed, since intermediate forms occur.
from the true N. vaiichcri in all of the fea- Other differences given by Argaman, such
tures listed by him as important in species as the shape of the flagellomeres, depend
differentiation, and, since the specimen he on the angle at which the specimen is
considered to be N. vimchcri was obtained viewed. Loss of the second submarginal
from Nonveiller, it is clear that Nonveiller cell (IS), thought characteristic of Ticopla
(1973) also misidentified the species. (Of by Brothers (1975), is also not significant
four specimens now in the Paris Museum since different degrees of reduction in
identified as N. viiuclicri by Andre, only wing venation are evident. We thus have
one (from Gibraltar, obtained from Saun- no hesitation in regarding Nauoniiitilla and
ders and thus almost certainly collected at Ticopla as synonymous. Lelej and Krom-
the same time as the specimen in the Lon- bein (2001) also regarded Ticopla as a syn-
don Museum) is correctly identified; the onym of Nauoniiitilla, implying that they
316 Journalof Hymenoptera Research
were following Argaman (1988) in this by Nonveiller (pers. comni.) before his re-
(which was incorrect since Argaman clear- cent death. The genus exhibits consider-
ly stated that he considered them distinct, able morphological variation (it is the only
see above); it is probable that they were ticopline genus in which brachypterous
really following the suggestion made by and apterous males are known) and is
Mitchell and Brothers (1998). The genus widely distributed throughout the Afro-
has an extensive distribution in the Afro- tropical, southern Palaearctic (Spain,
tropical and southern Palaearctic (Iberian North Africa) and Oriental Regions.
peninsula, Morocco, Algeria, Levant) Re- After this paper had been accepted for
gions, most species being as yet unde- publication, Lelej and Krombein (2001) de-
scribed. scribed three new genera of Smicromyr-
millini (CaiucroniUn, EosmicromifrmiUa and
Areotilla Bischoff
HiiuiiistniiiUn) from the Oriental Region
(Figs. 10-17)
and provided a key for their recognition.
Areotilla Bischoff 1920: 25, 174. For our study we had examined an apter-
Type species: Arcotilln areolahi Bischoff 1920, ous male of one of these genera {Hiiidus-
Transvaal, by original designation. tniiilln) and considered it to be a Siiiicw-
This genus was based on the type spe- lui/niiilln, although a somewhat anoma-
cies and A. mnrsluilli (Andre 1903). It is the lous one. We had also examined several
smallest genus in the subfamily in terms Afrotropical species with characteristics
of species numbers, and has been revised different from those Lelej and Krombein
by Mitchell and Brothers (1998). It com- considered limited to Suiicronn/nuilln, but
prises eight species known froni males again did not consider them as generically
and two known from females, all from distinct. We recognised that Suiicnvin/nuil-
southern Africa. (It should be noted that la, as we conceived it, was quite variable,
the correct spelling of the name for the but saw independent variation in several
new species referred to as A. fcrrugiiica (in of the characters used by Lelej and Krom-
the text) and A. fcrrugiiuitn (in a figure cap- bein (2001) te^ distinguish their new gen-
tion) by Mitchell and Brothers (1998) era, with many interniediates making rec-
should be A. ferniginen; the hip^sus is re- ognition of new putative genera question-
gretted.) able. For this reason, we do not distin-
guish between Sniicwiin/niiilhi and the
Sniicroiui/rntilla Suarez new genera proposed by Lelej and Krom-
(Figs. 18-25)
bein (2001), but do not wish to synony-
Smkwmyrmilla Suarez 1965: 570. mise them formally. As far as we can as-
Type species: Miitilla ariasi Andre 1896, certain, those genera agree with Siiiicw-
Spain, by original designation. nn/rniilln in all of the characters we have
This genus was described for a single used in this analysis. (We also suspect,
species (and single female specimen), al- however, that Camcrouilla may not actu-
though a second species from Spain, Suii- ally be a ticopline. Lelej and Krombein
cronn/rniilln nilrniuin Nonveiller and Gros based their conclusions entirely on the
1996, based on a single male specimen, rather inadequate original description and
has since been described; these are the figure of the female of Mutilla ocdipus
only specimens recorded from that coun- Cameron 1897 in placing it in this subfam-
try, and may thus be conspecific. Never- ily, citing the presence of a median and
theless, many other species of Sniicroiui/r- lateral spines on the propodeum. Those
niilla have been described and even more characteristics would not preclude its
await description; they were being revised placement in Myrmillinae, however.
Volume 11, Number 2, 2002 317
Figs. 10-17. Arcotilln spp. 10-13, A. fcrni;^iiicii Mitchell and Brothers, 9, length = 4.9 mm (paratype. South
Africa, KwaZulu-Natal). 14, 16, A. niarslmlli (Ancire), 6, length = 7.7 mm (holotype, South Africa, Northern
Province). 15, 17, A. vulgnris Mitchell and Brothers, 6, length = 8.9 mm (paratype. South Africa, Eastern
Cape). Scales = 1.0 mm.
318 Journal of Hymhnopthra Research
Figs. 18-25. Siiiicwiin/ri)tiUa spp. 18-21, S. sp., 9, length = 8.4 mm (South Africa, Mpumdliingd). 22-23, S.
sp., S, length = 6.5 mm (South Africa, Gauteng). Scales = 1.0 mm.
which placement is also suggested by its males, 8 on females) were surveyed. Of
enlarged quadrate head.) the more than 40 morphological characters
MATERIALS AND METHODS examined, 21 (Appendix 1) had appropri-
ate levels of variation (i.e. they were found
Specimens of all available species of Ti- to be variable among but not within gen-
coplinae {Arcotilhr. 8 species based on era) and were used in cladistic analyses.
niales, 2 ori females; Niiiioiinitilhr. 18 on Character polarity was established by out-
males, 8 on females; Siiilcwnn/niiilhr. 21 on group comparison. When there was vari-
Volume 11, Numbhr 1, 2002 319
Table 1. Data matrix for analysis of genera of Ticoplinae using 21 characters of Appendix 1.
320 Journal of Hymenoptera Research
0.62. Under fast (accelerated transforma- metasomal tergum. Macropterous males
tion) optimisation, the 23 derived states with fully articulating meso-metapleural
comprise 14 autapomorphies, 5 unique suture (i.e., no ventral bridge-like fusion
synapomorphies for Areotilla and Nauo- between meso- and metapleuron), poste-
mutilln, 1 unique synapomorphy for all riorly convex mesopleural margin, petio-
three genera, and 3 homoplasies (charac- late second submarginal cell in the fore-
ters 7M, lOM and 16M). An alternative wing, and volsella lacking digitus (i.e.,
tree with Areotilla and Suikrounjrnulln as only cuspis present). Females and microp-
sister groups (supported by 3 synapomor- terous/apterous males with mesosoma
phies) is two steps longer and has a much widest posteriorly (seldom with sides
lower RI (length = 28, CI = 0.82, RI = more or less parallel) with one or more
0.37); the other alternative tree with Na- weak to strong teeth or spines at postero-
noimitilla and Stiiicroiuynnilln as sister lateral angle, posterolateral margin of
groups is not supported by any synapo- pronotum indistinguishably fused with
morphies and is even longer (length = 31, mesopleuron (except in males with pron-
CI = 0.74, RI = 0.00). otum articulating with mesothorax), and
Froni the results, it is evident that Ar- distance from hunieral angle to pronotal
eotilla and Naiioiiiiitilla are the most closely spiracle at least as long as that between
related cladistically, and are distinct from pronotal and propodeal spiracles (except
Sniicronn/niiilln in several respects. The de- in males with articulating pronotum
gree of difference seen between the two where it may be shorter).
groups is similar to, if not greater than, Ticoplini Na—gy 1970 (= Nanomutillini
that between the tribes of Sphaeropthal- Suarez 1975). Eye strongly pubescent;
minae or Mutillinae (Brothers 1975) or antennal tubercles closely approximated
those of Myrmosinae (Brothers 1999). This basally but separate, not joined by a
contrasts with the opinion of Brothers straight transverse ridge, scarcelyprotrud-
(1975) who had seen far fewer represen- ing; pronotuni smoothly and evenly con-
tatives of these genera and who felt that vex over anterior declivity, without a
there were no marked groupings between transverse carina; propodeum with disc
them. We thus consider that recognition of and declivity distinct; second metasomal
two tribes, as proposed by Argaman sternum without felt line. Males with no-
(1988), is warranted despite the fact that tauli usually distinct (often faint and
both contain relatively few species when sonietimes absent in NtDiouiutilln); scutel-
compared with most other tribes of Mu- lum not apically produced; propodeum
tillidae. with three or five large fields covering en-
tire surface ofdisc and defined by well de-
CHARACTERISTICS OF SUBFAMILY
AND veloped carinae; metasomal sternum 2
TRIBES
with a short median longitudinal carina
Previous descriptions or diagnoses of basally; penis valve > 0.75X as long as
the subfamily and tribes, such as those by paramere. Females with at most one short
Brothers (1975, 1993), Suarez (1975) and spine on posterolateral angle of propo-
Argaman (1988), are incomplete or inac- deuni at apex of lateral oblique transverse
curate, mainly because those authors had carina; 'auricle' at base of first metasomal
access to far fewer species than we were tergum merely a small rounded protuber-
able to examine. The following descrip- ance; no defined pygidial area.
—
tions are followed by comments indicating Smicromyrmillini Argaman 1988. Eye
differences from previous attempts. pubescence absent although pores and/or
Ticoplinae Nagy 1970 (= Nanomutilli- very sparse minute setae may be present;
—
nae Suarez 1975). No felt line on second antennal tubercles fused basally,joined by
X
Volume 11, Number 2, 2002 321
a small straight transverse ridge, distinctly Argaman (1988) also included only Na-
protruding; pronotal dorsum sharply sep- nomutilla (and 'Ticopla'), but both sexes, in
arated from anterior declivity, with junc- the Ticoplini. He thought that the flagel-
tion angular and marked by a transverse lomeres were different in shape and struc-
carina; propodeum with disc and declivity ture from those in Smicromyrmillini, that
evenly merging, not distinct; second me- the pronotum (in the female) differed in
tasomal sternum with well developed lat- the number of lateral pits and that the de-
eral felt line. Macropterous males lacking gree of production of the apex of the pro-
notauli; scutellum apically produced over podeum at the articulation with the me-
metanotum; propodeum with threepoorly tasoma differed. We have found that fla-
defined anterior fields and many reticu- gellomere shape varies across both tribes
lations forming mini fields over posterior and also according to viewpoint, that the
half; metasomal sternum 2 lacking median development of pits on the pronotum
longitudinal carina; penis valve < 0.60 varies considerably and that the produc-
as long as paramere. Females and microp- tion of the propodeal lobe also varies and
terous/apterous males with at least two is not significantly different from the con-
spines on posterolateral angle of propo- dition in most other Mutillidae. Argaman
deum, lacking lateral oblique carina to remarked on the fact that the second me-
base of spine; 'auricle' at base of first me- tasomal tergum is longer than wide in fe-
tasomal tergum forming a strong lamel- male Ticoplini, but this is true only of
late or spinose protuberance; glabrous py- some Na)ioniutilla and not of Areotilla. For
gidial area well defined. male Ticoplini, Argaman noted the pres-
Suarez (1975) included only Nanomutilla ence of a single complete ridge on the
(females) in his subfamily Nanomutilli- scape, but this is true of Nanomutilla only
nae. Various of the features that he high- (see our character 7, below).
lighted as being characteristic of the group Neither Suarez (1975) nor Argaman
(as compared with Sniicroun/rniiUa which (1988) mentioned the pubescence of the
he placed in the Myrmillinae) are thus re- eye in female Ticoplini, presumably be-
stricted to that genus, and in particular to cause it is difficult to see in Nanomutilla
a species which he considered to be N. specimens, which are very small. Both au-
vnuchcri. Non-differentiated pubescence thors noted the absence of 'auricles'
on the body was thought characteristic, as (Brothers 1975) at the base of the metaso-
compared with the varied pubescence ma, but they are actually present although
generally forming patterns in other Mutil- inconspicuous.
lidae; although this is particularly true of For the Smicromyrmillini, Argaman
Naiioiiiiitilla, it is approached in Areotilla (1988) made much of the carinate anterior
but is not particularly significant since it margin of the mesoscutum in the male,
occurs elsewhere in the Mutillidae also. considering this a unique character in Hy-
The peculiar 'bethyloid' or 'proctotrupoid' menoptera; such a carina is certainly pre-
bodv form was also highlighted, but this sent in some species of Smicromi/rmilla but
is not true of Areotilla. many other species have no trace of it.
KEY TO TRIBES AND GENERA OF TICOPLINAE
[Note: We consider Eosmicnvm/rmilla and Hindustanilla doubtfully distinct from Smicwmynmlla; Camcwnilla is
probably misplaced in Ticoplinae, see above, and is therefore omitted from this key.]
1 (a) Wings well developed (male) 2
(b) Wings absent or rudimentary (female, rarely male) 4