The monotypic family Carthaeidae Common, 1966 includes only a single species, Carthaea saturnioides Walker, 1858, and is endemic to Australia. Adults of this large species have a very beautiful wing pattern with very prominent eye-spots, making them a rather popular icon for nature-related publications and web pages. Morphologically, the species stands out from other Bombycoidea by its retention of several symplesiomorphies (shared, ancestral characteristics), which is why the family was considered to be a particularly "primitive" member of the superfamily Bombycoidea (Common 1966). Little is known about the species beyond the thorough descriptions by Common (1966, 1990; largely morphology) and the illustration of eggs and larval records by McFarland (1973, 1979). Although specimens are present in the Australian National Insect Collection (Canberra) and larvae have been raised from eggs by a few lepidopterists, nothing has been published on the larval stages preceding the penultimate instar described by Common (1966). Likewise, published life history observations are limited to Common (1966) and McFarland (1979). This lack of information is largely due to the very limited distribution of the species.
This large (76-100mm wingspan), magnificent moth is easily recognized by its blue eye-spots on the hindwings and the pair of bright white bands on the forewing. Females resemble males in appearance, but have thin, serrate antennae (rather than strongly bipectinate antennae) and are larger in size. Larvae have a similarly striking appearance with prominent lateral eye-spots on all abdominal segments. These larval eye-spots and dorsal, transverse rows of deep pits on the pupal abdomen are unique, derived characteristics (apomorphies) of the species (and, hence, family). However, the species was primarily placed in a family of its own (Common 1966) due to the presence of several "ancestral" (symplesiomorphic) characteristics that are rarely found within the superfamily Bombycoidea: three-segmented maxillary palpi; well developed proboscis with carinate papillae; frenulum and retinaculum in both sexes; completeness of the radius in the forewing. Not fitting into any of the other bombycoid families and including only a single species with very limited distribution, the family Carthaeidae is often considered as isolated within the Bombycoidea and was regarded as a particularly "primitive" member of the Bombycoidea (Common 1966).
Geographic Range description:
Endemic to the southern part of Western Australia, ranging south from Northampton and as far east as Mt Ragged (Cape Arid National Park) (Common 1990).
The species (and, hence, the family) is endemic to the southern part of Western Australia, ranging south from Northampton and as far east as Mt Ragged (Cape Arid National Park) (Common 1990). Throughout this distribution, the species is rather rarely encountered, but can be locally common in undisturbed areas.
Pale orange-yellow (Common 1990).
Egg mass pattern:
Attached to the young food plants in groups of two or three (Mcfarland 1973).
Description of egg morphology:
Flat type, broadly oval, with micropyle at one end; chorion smooth and somewhat shiny, not as tough as in other Bombycoidea (Common 1990).
Head hypognathous, with the frontoclypeus extending less than half the distance to the epicranial notch (Common 1990). Head capsule with a dark markings that change with instars. Mandibles without secondary setae (Common 1966).
Body setae on verrucae:
Body setae on chalazae:
Body setae on scoli:
Larval body description:
Secondary setae minute (Common 1990); first to penultimate instar with scoli that carry primary setae (black with white, clubbed apex in at least penultimate instar) and short secondary setae; scoli and setae strongly reduced to lost in final instar (Common 1966, 1990). Detailed description of penultimate and final instar in Common 1966.
Larval thorax description:
Meso- and metathorax with D1 and D2 on single large scolus (penultimate instar; Common 1966).
Larval abdomen description:
Segments A1 to A8 with prominent lateral eye-spots surrounding spiracles (Common 1966, 1990); A8 with D1 scoli merged, forming a middorsal horn (up to penultimate instar, reduced to hump in final instar; Common 1966, 1990).
Crochet arrangement description:
Penultimate and final instar with simple, biordinal crochets arranged in meso-series (Common 1966, 1990).
Anal comb on A10:
Strongly sclerotized, dull blackish; head, wings and appendages rugose; abdomen pitted, especially on dorsum where there is a anterior row of deeper pits on segments 1-8; eyes small, maxillary palpi indicated by small triangular sclerites lateral of the eye; antennae strongly bipectinate to apex in male, extending nearly to tips of wings; haustellum slightly exceeding tips of wings; abdominal segments 5 and 6 movable, with visible ventral proleg scars; abdominal segment 10 deeply excavated ventrally, produced and tapering posteriorly, forming a cremaster with a series of hooked setae (Common 1966).
Pupal tergal spines:
A very flimsy and loose cocoon is spun on the ground, underneath debris (observation in captivity; pupation occurs soon after spinning of the cocoon (Common 1966).
Adult Abdomen Morphology
Female genitalia description:
Papillae anales with long hairs; apophyses posteriores fairly short and slender; apophyses anteriores forming strong Y in basal half; ostium bursae very broad and shallow; lamella antevaginalis in form of a broad strongly sclerotized pocket with broad posterior opening, containing dense scales; bursa copulatrix very short; ductus bursae short and broad; ductus seminalis leaving ductus bursae close to ostium bursae; corpus bursae a small sac with thickened walls; no signum (Common 1966). Joined reservoirs of the two accessory glands forms a common duct that is set off and at right angle to the joined reservoirs.
Female pregenital sexual scales:
Female accessory glands:
Female oviduct opening:
Female bursa ostium opening:
between S7 and venter 8
Female anterior apophyses originating:
originating from T8, from venter 8
Male pregenital sexual scales:
Male genitalia description:
Tegumen broad and well sclerotized; uncus tapering posteriorly, apically blunt; gnathos with sclerotized arms, forming a broad median plate covered with minute spines; possibly secondary sclerotization between gnathos and valvae reduced; juxta simple; vinculum ventrally narrow and rounded; valva large, rounded and only moderately sclerotized; phallus cylindrical, strongly ventrad curved and with a coecum penis; vesica with strongly sclerotized, apically dentate cornutus (Common 1966, 1990; Zwick 2009).
Sternum 5 gland:
Adult abdomen description:
Stout and clothed in long, dense piliform scales (Common 1966); lateral bars of dorsum 1 strongly produced ventrad (posteriorly), coming very close to sternum 2; no venulae; sternum 2 with short apodemes and, in males, with androconial tufts attached to sclerotized arms (Lemaire and Minet 1998).
phallotheca and aedeagus (phallus)
Adult Thorax Morphology
Adult thorax description:
Densely clothed in piliform scales (Common 1966, 1990); spinarea absent (Common 1990; Lemaire and Minet 1998).
Clothed with long piliform scales; fore leg with a strong epiphysis in both sexes, nearly reaching to the distal end of the tibia; tibial spurs number 0-2-4 and with appressed scales and scattered short, stout setae, but naked spur apices; inner tibial spurs longer than outer ones (Common 1966, 1990); tarsi longer than corresponding tibiae and densely bristled beneath; arolium well developed; pulvilli with long, dorsal setiform outgrowths; pretarsal claws simple (Common 1966, 1990; Lemaire and Minet 1998).
Forewing discal cell about half the length of the wing; R free; Rs complete (Rs1-4), with Rs1+Rs2 long-stalked, but Rs3+Rs4 short-stalked; M1 at upper angle of cell; M2 at middle of cell; M3 at lower angle of cell; CuA1 and CuA2 well before the lower angle of the cell; CuP absent; 1A+2A form a small basal fork (Common 1966, 1990; Lemaire and Minet 1998). Hindwing without expanded humeral area and humeral veins; Sc separate from R at the base, but joined by R at about basal quarter of discal cell, with Sc+R gradually diverging from Rs; peripheral veins arise separately from the cell, with Rs arising well before upper angle of cell and M2 being slightly closer to M1 than to M3; CuP absent; 1A and 2A separate; 3A present (Common 1966, 1990).
Forewing cell veins:
Forewing anal vein notation:
1A+2A forming a narrow basal fork.
Forewing upper surface with microtrichia:
Hindwing anal vein notation:
1A reaching tornus; 2A vestigial except near base; 3A rather short (Common 1966, 1990; Lemaire and Minet 1998).
Hindwing cell vein:
Wingspan 76-100mm; forewings broad, triangular and with a prominent eye spot in both sexes; retinaculum membranous in males, absent in females (Common 1990).
Hindwings broad and with a prominent eye spot in both sexes; frenulum a strong bristle in males, but in females a group of about 10 short, fine bristles, which are probably functionless (Common 1990).
Adult Head Morphology
Labial palpus modification:
Prominent, 3-segmented, reaching to near base of antennae; second segment ascending, third segment porrect or ascending; basal two segments thickened with dense long hairscales, apical segment slender with appressed scales (Common 1966).
Fluted sensilla styloconia on proboscis:
Well developed and coiled, with carinate papillae [= fluted sensilla styloconia] towards apex (Common 1966).
Head vertex scaling:
Female flagellomere description:
Axis of flagellum scaled dorsally to apex (Common 1966).
Male flagellomere description:
Axis of flagellum scaled dorsally to apex (Common 1966). Bipectinate to apex with long rami, and additionally with a well developed, ventro-median sensory process ("tri-pectinate").
General antennae description:
Longer than thorax.
Adult head description:
Head clothed in piliform scales; chaetosemata absent; pilifers well developed, with dense bristles; maxillary palpi very small and three-segmented, basal segment bearing a proximal seta, second and apical segments with long scales, the latter with short apical projections (Common 1966, 1990; Lemaire and Minet 1998).
Larva with prominent eye-spots surrounding the abdominal spiracles (Lemaire and Minet 1998). Larva with prominent eye-spots surrounding the abdominal spiracles (Lemaire and Minet 1998)
Comparison with similar species
While the species name, C. saturnioides, hints at the superficial similarity with emperor moths (Saturniidae) due to the very prominent eye-spots, the species has a very distinct and unique appearance and can't be confused with any other known species.
The species occurs from coastal, arid scrubs to tall forests, depending on the availability of its host plants.
Life History and Behavior
Resting adults hold their wings in a roof-like position, but display a defence behaviour if disturbed. Pressing head and posterior end of the abdomen against the ground, the forewings are opened and inclined forward, revealing the colourful hindwings with the large, prominent eyespots. While remaining in this posture, the hindwings are rhythmically moved sideways at a rather slow pace (Common 1966). Males are readily attracted to light past 10pm, while females are more often encountered in flight between low vegetation rather than at light. Males held with their wings closed over the abdomen will oftentimes display their androconial tuft of scales on A2.
Life History: Immature Stages
Pupal stage lasts from summer to spring.
Larval food items include:
Dryandra cirsioides (Proteaceae) (Common 1966). Dryandra hewardiana (Proteaceae) (Common 1966). Dryandra pteridifolia (Proteaceae) (Common 1990). Dryandra fraseri (Proteaceae) (Common 1990). Banksia sphaerocarpa (Proteaceae) (Common 1990). Banksia caleyi (Proteaceae) (Common 1990). Grevillea concinna (Proteaceae) (McFarland 1979; Common 1990)
Larval food habits description:
Larva feed exposed on young spring foliage (Common 1966).
Life history larvae:
Presumably five instars (Common 1966). Larva lethargic.
Life habits of immature stages:
The species has presumably five larval instars, of which only the last two have been described in publications so far (Common 1966, 1990). Up to the penultimate instar, larvae have numerous protrusions (scoli) that carry a few, thick hairs, some of which have clubbed ends; in contrast, the final instar has a smooth, "naked" appearance. Larvae hatch from rather flat, ovoid eggs in early spring and feed fully exposed on the young foliage of primarily Dryandra and Banksia (both Proteaceae). Pupation occurs at the beginning of summer in a very flimsy and loosely spun cocoon on the ground, and the pupal stage lasts through the hot summer as well as the winter, with adults emerging in spring.
Life History: Adults
While C. saturnioides has a very well developed proboscis and is likely to be a nectar feeder, actual feeding hasn't been observed so far.
Life history adult:
One generation per year, with adults being on the wing from October to December, even during cold, windy spring nights.
Life cycle description:
Larvae have five instars.
Life habits of adults:
While the adults have a very well developed proboscis, it is unknown if it is used for nectar feeding. Little is known about the life history of adults, except for a characteristic defence behaviour. If disturbed, the moth presses its head and tip of the abdomen against the ground, which inclines its opened forewings forward and reveals the colourful hindwings with the large, prominent eye-spots. While remaining in this posture, the moth rhythmically moves its hindwings from side to side at a rather slow pace (Common 1966).
Evolution and Systematics
This large (76-100mm wingspan), magnificent moth is easily recognized by its blue eye-spots on the hindwings and the pair of bright white bands on the forewing. Females resemble males in appearance, but have thin, serrate antennae (rather than strongly bipectinate antennae) and are larger in size. Larvae have a similarly striking appearance with prominent lateral eye-spots on all abdominal segments. These larval eye-spots and dorsal, transverse rows of deep pits on the pupal abdomen are unique, derived characteristics (apomorphies) of the species (and, hence, family). However, the species was primarily placed in a family of its own (Common 1966) due to the presence of several "ancestral" characteristics (symplesiomorphies) that are rarely found within the superfamily Bombycoidea: three-segmented maxillary palpi; well developed proboscis with carinate papillae; frenulum and retinaculum in both sexes; completeness of the radius in the forewing. Not fitting into any of the other bombycoid families and including only a single species with very limited distribution, the family Carthaeidae is often considered as isolated within the Bombycoidea and was regarded as a particularly "primitive" member of the Bombycoidea (Common 1966).
Systematic and taxonomic history
The very distinct and beautiful species Carthaea saturnioides had been described in the family Ommatophoridae [= Noctuidae] by Walker (1855). Subsequently, it was included in the Saturniidae (Kirby 1892) and the Geometridae (Prout 1910, 1912 and 1920; Turner 1930), until Common (1966) placed the species in a family of its own within the Bombycoidea. Based on the presence of several symplesiomorphies that have been lost in many of the other bombycoid families (e.g., three-segmented maxillary palpi; well developed proboscis with carinate papillae; frenulum and retinaculum in both sexes; completeness of the radius in the forewing), Common (1966) assumed the family to be "possibly the most primitive family of the Bombycoidea". Based on morphological characteristics, Minet (1994) proposed a phylogeny of the Bombycoidea, in which the Carthaeidae are the sister to the Sphingidae, Brahmaeidae and Lemoniidae [= Brahmaeidae], and all of these together the sister to the remaining Bombycoidea sensu Minet 1994 (Eupterotidae, Saturniidae, Bombycidae, Endromidae and Mirinidae). In contrast, the analyses of DNA sequences by Regier et al. (2008) and Zwick (2008) both place the Carthaeidae in a monophylum comprising the Anthelidae, Endromidae, Mirinidae and several subfamilies of the polyphyletic Bombycidae. In particular, the results of Regier et al (2008) provide strong support for this monophylum (bootstrap support of 97), but relationships within this group are essentially unresolved. Unpublished results from the LepTree project strongly support a sistergroup relationship between the Australian Carthaeidae and the Neotropical bombycid subfamily Phiditiinae.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:20
Specimens with Barcodes:19
Species With Barcodes:1
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