Holotype : NEW ZEALAND • ♂; Mid Canterbury [MC], Christchurch Botanic Gardens; 43°31.8'S, 172°37.2'E; emg. 21 Apr. 2014; R.J.B. Hoare, B.H. Patrick leg.; larva in leaf-fold on Teucridium parvifolium 31 Mar. 2014; NZAC.

Paratypes : NEW ZEALAND • 1 ♀; same collection data as holotype; emg. 24 Apr. 2014; NZAC • 3 ♂♂; MC, Christchurch Botanic Gardens; 31 Mar. 2014; R.J.B. Hoare, B.H. Patrick leg.; beaten from Teucrium parvifolium [as Teucridium on labels]; NZAC • 2 ♀♀; same collecting data as preceding; ♀ genitalia on slides NZAC Grac. 2, NZAC Grac. 4; NZAC • 4 ♂♂; MC, Christchurch, Kennedy’s Bush Rd, Jensen property; 29 Mar. 2014; R.J.B. Hoare, B.H. Patrick leg.; on and around Teucrium parvifolium; ♂ genitalia and wings on slide NZAC Grac. 3; NZAC • 1 ♂; MC, Banks Peninsula, Prices Valley; 1 Apr. 2014, R.J.B. Hoare, B.H. Patrick leg.; beaten from Teucrium parvifolium; ♂ genitalia on slide NZAC Grac. 1; NZAC.

Sabulopteryx botanica is distinctive amongst New Zealand gracillariids in its combination of small size (wingspan 10 mm or less) and yellow-ochreous black-speckled forewings. It is perhaps most similar to Caloptilia selenitis (Meyrick, 1909), but this species has the centre of the vertex white and has three white spots along the forewing dorsum (there is no white on the vertex or forewing in S. botanica). In the male genitalia, the paired processes on the dorsum of the valva are diagnostic, and in the female, the deep invaginations of the S7–S8 intersegmental membrane are characteristic.

Wingspan 8.5–10 mm. Adult male (Fig. 2): Head: frons white; vertex yellow-ochreous with some scales tipped darker brown; labial palpus whitish with segments 2 and 3 tipped brown; antenna ochreous, ringed dark brownish (apex of each flagellomere), approximately equal in length to forewing; scape with inconspicuous pecten of ca 5 short bristles (often abraded away). Thorax yellow ochreous with tegulae anteriorly blackish. Forewing: yellow-ochreous, with numerous blackish scales forming variable and irregular pattern of broken transverse fasciae; blackish scales often denser towards base of costa and in disc at ca 2/3 length of wing; fringe ochreous whitish, darker around apex to tornus, where dark-tipped scales form three indistinct fringe-lines (in fresh specimens). Hindwing pale greyish; fringe greyish white. Underside: forewing dark brown, paler on dorsum below fold, yellowish around base of cilia; hindwing dark brown on costa and dorsum, pale greyish centrally. Legs: foreleg and midleg with femur and tibia thickened with blackish scales and tarsi yellowish, each tarsomere tipped blackish above; hindleg yellowish, femur with black central patch exteriorly, tibia ringed brownish subapically and each tarsomere with a few brownish apical scales. Abdomen silvery grey, with yellowish anal tuft.

Figures 2, 3. 

Sabulopteryx botanica, adults. 1 Male paratype, Halswell Quarry (Kennedy’s Bush Rd), Christchurch MC, 29 Mar 2014 (NZAC) 2 female paratype, Christchurch Botanic Gardens MC, emg. 24 Apr 2014 (NZAC).

Adult female (Fig. 3). As described for male, but abdomen tipped with glossy ochreous whitish scales.

Wing venation (Fig. 4). Forewing 12-veined, as described for the genus by Triberti (1985), who regarded the 12-veined condition as being due to coincidence of M2 and M3. Discal cell somewhat dilated posteriorly as described by Triberti (1985). Hindwing very narrow (more so than in other Sabulopteryx species) with Rs strongly approximated to costa for most of its length; cell open between M2 and M3; M3 and CuA1 closely approximated and parallel.

Figures 4–6. 

Sabulopteryx botanica, adult morphology. 4 Wing venation 5 male abdominal base 6 male abdomen, segments 7–8 (sternites on left).

Male abdomen and genitalia (Figs 5–8). Abdominal base as in Fig. 5. S7 and S8 (Fig. 6) markedly shortened and much wider than long, each with lateral pair of coremata. T8 (Fig. 6) with T-shaped sclerite. Genital capsule (Fig. 7): tegumen rather weakly sclerotised, elongate-triangular with attenuate apex. Tuba analis longer than tegumen; subscaphium slender, weakly sclerotised. Valva oblong, narrowed at base, with rounded apex; apical third bearing numerous long fine setae directed obliquely towards costa. Base of valva complex: costa extended into anteriorly-directed narrow, weakly curved process dorsad of anellus membrane (not fused with process from opposite valva, i.e. forming transtilla broken in the middle); from base of this process sclerotised ridge extends across inner (ventral) face of valva to base of long, sclerotised weakly curved spine that extends from valval dorsum at 1/3 valva length; a second, similar spine (slightly more strongly curved) on valval dorsum at 1/2 valva length. Juxta absent. Vinculum large, oval, saccus not differentiated. Phallus (Fig. 8) very elongated, slender, with sharply pointed apex; basally extending smoothly into ovoid bulbus ejaculatorius; vesica without cornuti; caecum penis absent.

Figures 7–11. 

Sabulopteryx botanica, adult and pupal morphology. 7 Male genital capsule 8 male phallus 9 female genitalia 10 pupal exuviae, head, ventral view 11 pupal exuviae, thorax and abdomen, dorsal view.

Female genitalia (Fig. 9). Ovipositor lobes rather short, membranous, basally with long setae, setae denser and shorter towards apex of each lobe; apophyses posteriores short. S7-S8 intersegmental membrane with pair of deep membranous sublateral invaginations; ostium lying between these, dorsal wall of ostium extended into T-shaped membranous area bordered posteriorly by strongly sclerotised transverse lamella postvaginalis, which is continuous with and broadens into lateral sclerotisations of S8. Ductus bursae entirely membranous, long and slender, ca 3× length of corpus bursae; corpus bursae more or less ovoid, with pair of spine-like signa, one long and one very short.

Immature stages. Egg. Elongate-oval, flat, showing up as silvery white translucent ‘shell’ at start of mine, apparently with rather coarsely sculptured chorion (not observed under SEM). Larva (Fig. 12). Head translucent pale yellow-brown, margined dark brown posteriorly and along adfrontal / ecdysial lines; blackish in region of stemmata. Body translucent yellowish white, with the gut contents showing through bright green; prothoracic plate in form of two irregular r-shaped sclerites with outlying smaller sclerites anteriorly. Thoracic legs with sclerotised areas dark grey-brown. Prolegs present on A3–5 and A10; crochets on A3–5 biserial: outer row a complete circle, with anterior crochets reduced, inner row a transverse semicircular band of larger crochets in posterior half of planta; A10 with single transverse band of large crochets in anterior half of planta. Anal plate a small transverse brownish sclerite with poorly defined margins. Chaetotaxy as described and figured for Aspilapteryx tringipennella (Zeller, 1839) by Triberti (1985). Pupa. Head (Fig. 10): frons smoothly rounded, without processes, without setae near antennal bases; antennae ca 3× as long as labial palpi. Thorax (Fig. 11): mesothorax and metathorax each with one pair of well-developed dorsal setae; forewings reaching to ca 1/2 way along A5, hindwings to A3/A4 junction. Abdomen (Fig. 11): A2–8 each with irregular rows of spinules dorsally, spinules slightly smaller and more widely spaced on A2; A7 not furrowed ventrally; abdominal tip truncate, with 3 pairs of small spinose tubercles.

Figures 12–15. 

Sabulopteryx botanica, larva, host plant, and early mine. 12 Larva, dorsal 13 Teucrium parvifolium in flower, Longbush, Wairarapa (photo courtesy of J. Rolfe) 14 initial linear mine on leaf underside 15 early blotch mine.

The only known host plant is the small-leaved divaricating shrub Teucrium parvifolium (Lamiaceae) (Fig. 13), which is endemic to New Zealand and was until recently placed in its own monotypic genus Teucridium, and in the family Verbenaceae. Salmaki et al. (2016) showed that Teucridium belongs to Lamiaceae and is phylogenetically nested within the large worldwide genus Teucrium. The shrub is widespread on both main islands of New Zealand, but rare and very local, and has a conservation status of ‘At Risk – Declining’ (de Lange et al. 2018). The habitat is described as ‘fertile stream sides and river terraces in lowland dry forest and podocarp-hardwood forest, occasionally on forest margins, clearings and amongst scrub’ (NZPCN 2019). These fertile alluvial habitats have been cleared of forest throughout much of the country. Since no other species of Teucrium is native to New Zealand, Sabulopteryx botanica must be considered strictly monophagous on T. parvifolium (see also Remarks below).

The egg is laid on a leaf of the host plant, usually on the underside next to the midrib. The young sap-feeding larva forms a linear white mine (Fig. 14), almost invariably on the leaf underside, that extends to the margin of the leaf (on the side of the midrib that the egg was laid), then typically crosses the midrib at the leaf apex and extends for some way down the leaf margin on the other side. A line of blackish frass is more or less visible in the centre of the mine at this stage. The larva then doubles back and begins to expand the mine into a white blotch (Fig. 15), usually concentrated towards the leaf apex or to one side of the midrib, but often taking up the whole leaf on smaller leaves. These early mine stages are often rather hard to see unless the leaf is examined closely from the underside. Occasionally the egg and initial mine are on the upperside. Later the larva expands the mine and spins silk extensively in the interior (on the eroded inner surface of the leaf underside), causing the leaf to fold and creating creases in the underside (Fig. 16), in the typical manner of many gracillariid miners. At this stage, patches of the palisade mesophyll are eaten, leaving small windows of upper epidermis towards the middle of the leaf (appearing like holes), and larger windows (browning with age) towards the leaf margin. Black frass is scattered across the inner surface of the upperside of the leaf. When full-fed, the larva leaves the mine and folds a fresh leaf in half from the underside (Fig. 17), forming a cocoon of dense white silk within, in which it pupates. (In captivity, some larvae spin cocoons in tissue paper at the bottom of the rearing container.)

Figures 16–18. 

Sabulopteryx botanica, late mine, cocoon, and parasitoid. 16 Fully formed mine with silk spinning causing creases 17 folded leaf with cocoon 18 Sympiesis sp. (Hymenoptera: Eulophidae) partially emerged from cocoon of S. botanica in leaf from herbarium sheet AK285999 (Auckland Museum).

One species of hymenopteran parasitoid can be confidently associated with the early stages of S. botanica, and one tentatively. A specimen of an unidentified species of Sympiesis Förster, 1856 (Eulophidae: Eulophinae) was found partially emerged from a cocoon in a folded leaf on an Auckland Museum herbarium sheet (Fig. 18). The plant specimen (AK285999) was collected at Pareora Scenic Reserve SC on 17 Mar 2004 by P.J. de Lange. (The wasp specimen was removed and mounted, and is now in NZAC, cross-referenced with the herbarium sheet.) Interestingly, two species of Sympiesis (S. euspilapterygis (Erdös, 1958) and S. gregori Boucek, 1959) have been associated with the Teucrium-mining Sabulopteryx limosella in Europe, but both also attack other leaf-mining Lepidoptera (see references in Noyes 2018, De Prins and De Prins 2018).

One specimen of an unidentified species of Dolichogenidea Vierek, 1911 (Braconidae: Microgastrinae) was reared from amongst Teucrium leaf-mines collected in Christchurch Botanic Gardens on 23 Jan 2018, emerging on 29 Jan (NZAC). It is thought most likely that this wasp was a parasitoid of S. botanica; however, host remains were not found and the sample was discovered later to include one unidentified early instar tortricid larva (preserved, not parasitised). The genus Dolichogenidea does not appear to have been associated with Sabulopteryx before, but is recorded overseas from other Gracillariinae (Caloptilia spp. and Gracillaria syringella (Fabricius, 1794)) as well as Lithocolletinae (Phyllonorycter spp.), Ornixolinae (Parectopa ononidis (Zeller, 1839)) (De Prins and De Prins 2018) and many other Lepidoptera, especially microlepidoptera (Austin and Dangerfield 1992). Most reared material of New Zealand Sympiesis and Dolichogenidea spp. in NZAC is associated with larvae of Tortricidae (Tortricinae).

New Zealand, from the following regions: CL, TO, GB, HB, RI, WI, WA / NN, MC, SC, CO.

The adult moth has only so far been found or reared in mid Canterbury (MC), but records of leaf-mines and cocoons on herbarium specimens of the host reveal a much wider range (Fig. 19). In some of these areas the plant is very likely to be severely threatened or even extinct, and renewed searches for plant and moth are desirable throughout the country. Towards the north and south of the plant’s range, herbarium records of mines are scarce. The only Coromandel record is from a herbarium specimen collected at Kauaeranga near Thames prior to 1906 by J. Adams (Auckland Museum AK108237); no recent material of the plant from this area was seen. The only Otago record is from Gorge Creek, near Roxburgh CO, where P.N. Johnson found a colony of Teucrium in a shaded rock cleft on 24 May 1993 (Allan Herbarium CHR481347; two early mines and one cocoon). It should be noted that only two major herbaria were visited during the course of this research, and there are likely to be preserved mines in other botanical collections that have not yet been visited. The host plant is not known from any offshore islands of New Zealand, so these have been omitted from the map (Fig. 19).

Figure 19. 

Distribution of Teucrium parvifolium and Sabulopteryx botanica based mainly on records from herbarium sheets in Allan Herbarium, Lincoln and Auckland Museum herbarium.

Adults have been found in the wild or emerged from wild-collected larvae or pupae in every month of the year in mid Canterbury, and the species is probably more or less continuously brooded here. All stages from early mines to adults can usually be found in the Christchurch Botanic Gardens, where the species is common amongst its host. Phenology in other parts of the range is unknown.

The species name refers to the close association of this species with botany and botanists. It was discovered by a botanist (AE Esler) as a pressed larva in a botanical specimen of the host plant, and many further leaf-mines have now been found on herbarium sheets collected as part of botanical surveys. Its discovery by BHP as an adult in the type locality, Christchurch Botanic Gardens, completes the association.

Sabulopteryx botanica was accorded ‘Nationally Vulnerable’ status in the latest review of the conservation status of New Zealand Lepidoptera, where it was listed as Caloptilia sp. “Teucridium” (Hoare et al. 2017). This ranks as the third most critical category assigned to extant species (after ‘Nationally Critical’ and ‘Nationally Endangered’), and was based on the rarity and declining status of the moth’s host plant. As noted above, survey for S. botanica through most of the range of its host plant has been inadequate; most herbarium records of mines are over 25 years old (Fig. 19), and further field-work is needed to determine the moth’s current distribution. Teucrium parvifolium is an attractive, easily propagated and cultivated shrub that is tolerant of a wide range of conditions (NZPCN 2019), though relatively short-lived (P. Bellingham, pers. comm.). The moth appears to be thriving in situations where T. parvifolium has been planted around Christchurch, for example in the Botanical Gardens and in the native plantings maintained by botanist Carol Jensen at Kennedy’s Bush near Halswell Quarry. It should therefore be possible to boost the moth’s population substantially by encouraging the propagation and cultivation of the host plant, with due attention to appropriate sourcing and hygiene. This could be achieved in natural environments where the plant persists, as well as in public plantings and in suburban gardens.

The new species described here can be confidently placed in the Gracillaria group of genera (Gracillariinae) as defined by Kumata (1982), based especially on the following characters: mid femur and tibia thickened beneath with rough scales; R1 of forewing arising near base of wing, with upper vein of cell weakened beyond branching point of R1; hindwing R2+3 very short and running parallel with and very close to apical part of Sc+R1; hindwing cell open between M2 and M3; male segments 7 and 8 weakly membranous, with coremata. The following genera belong to the Gracillaria group, based on Kumata (1982) and updates from subsequent authors (e.g., Triberti 1985, Huemer et al. 2016, Pereira et al. 2019): Aspilapteryx, Caloptilia Hübner, 1825, Calybites Hübner, 1822, Ectropina Vári, 1961, Eucalybites Kumata, 1982, Euspilapteryx Stephens, 1835 (treated as a synonym of Calybites by Kumata (1982)), Gracillaria Haworth, 1828, Mercantouria Huemer, Lopez-Vaamonde & Triberti, 2016, Povolnya Kuznetzov, 1979, Sabulopteryx and Vallissiana Pereira & Arévolo, 2019.

When first discovered as an adult by BHP in January 2013, and before detailed morphological examination, S. botanica was tentatively considered to be a member of the genus Caloptilia (Hoare et al. 2017: see above). However, RJBH later noted its remarkable external similarity to some west Palaearctic gracillariids then placed in the genus Aspilapteryx (subgenus Sabulopteryx), i.e. S. limosella from central and southern Europe and S. inquinata from southern Europe, Turkey and Lebanon, which it closely resembles in size, wing shape and overall coloration. When describing Sabulopteryx as a new subgenus of Aspilapteryx, Triberti (1985) anticipated the possibility that it might deserve full genus status. In a recent molecular phylogeny, Pereira et al. (2019) found a 14 to 16% divergence in DNA barcodes between Aspilapteryx and Sabulopteryx species, and indicated that Aspilapteryx is polyphyletic if Sabulopteryx is included. Our analysis (see below) also retrieves Aspilapteryx and Sabulopteryx in separate lineages, so we accept the conclusions of Pereira et al. (2019) and treat Sabulopteryx as a genus.

As pointed out by Huemer et al. (2016), morphological comparisons within the Gracillaria group are complicated by the apparently homoplasious distribution of character states amongst genera. Based largely on comparison with the descriptions and figures in Vári (1961), Kumata (1982), Triberti (1985), Huemer et al. (2016) and Pereira et al. (2019), the characters listed below in combination lend support for placing the New Zealand Teucrium-miner in Sabulopteryx:

1. Male abdomen with coremata on both segment 7 and segment 8 (Fig. 6). This conforms with most genera of the Gracillaria group, including Sabulopteryx and Mercantouria (Huemer et al. 2016), but not with Gracillaria or Aspilapteryx, where there is only one pair of coremata (Kumata 1982; Triberti 1985), nor with Vallissiana, where there are no coremata (Pereira et al. 2019).

2. Outline of male valva (beyond sacculus) rounded, not angular, and lacking a ventro-apical lobe. In its rounded / oblong valva, S. botanica resembles most genera of the Gracillaria group, but not Mercantouria, Calybites or Euspilapteryx, all of which have a distinctly angular valva (Kumata 1982; Huemer et al. 2016), nor Aspilapteryx or Vallissiana, both of which have a distinct ventro-apical lobe (Pereira et al. 2019).

3. Setae of valva confined to apical area, not extending into basal half. This character does not appear to have been commented on by previous authors: Caloptilia, Gracillaria, Povolnya and Calybites all have the valva more extensively setose than the remaining genera of the Gracillaria group (including Sabulopteryx), perhaps as a result of the relative reduction of the (non-setose) sacculus in these four genera.

4. Valva lacking stout peg-like or spine-like setae distally. This conforms with most genera of the Gracillaria group, including Sabulopteryx. Short, stout setae are present in the distal part of the valva in Euspilapteryx and on the ventrodistal margin in Calybites (Kumata 1982), and longer, spine-like setae in Eucalybites (Kumata 1982) and Mercantouria (Huemer et al. 2016).

5. Male phallus short and straight, without apical processes. The phallus of S. botanica (Fig. 8) is similar to those of described species of Caloptilia, Gracillaria, Povolnya and Sabulopteryx. It lacks the apical modifications found in Eucalybites (Kumata 1982: figs 47 B, C), Euspilapteryx and Vallissiana (Pereira et al. 2019: fig. 3F) and the rod-like apical sclerite of Mercantouria (Huemer et al. 2016: fig. 5). The phallus is long with a helical tip in Aspilapteryx (Triberti 1985; Huemer et al. 2016), very long and straight in Calybites (Kumata 1982), and curved or sinuous in Ectropina (Vári 1961).

6. Female genitalia with two curved, spine-like signa (Fig. 9). This is typical of Aspilapteryx and Sabulopteryx (Triberti 1985), Mercantouria (Huemer et al. 2016), most Caloptilia and Eucalybites (Kumata 1982), but not of Gracillaria, Caloptilia subgenus Minyoptilia Kumata, 1982, Calybites, Ectropina, Euspilapteryx, or Vallissiana, in all of which there is only a single signum (Vári 1961; Kumata 1982; Pereira et al. 2019). Povolnya has two signa, but these are short and stout (Kumata 1982).

7. Forewing brownish, without costal streak and with numerous irregularly arranged darker blotches (Figs 1, 2). This wing pattern accords with the description of Sabulopteryx by Triberti (1985), and as noted above, there is a strong superficial resemblance between adults of S. botanica and the two Palaearctic members of Sabulopteryx, S. limosella and S. inquinata. No other member of the Gracillaria group closely approaches this wing pattern.

8. Host-plant genus Teucrium. The hostplant genus is shared with S. limosella, type species of Sabulopteryx, which mines in Teucrium chamaedrys L. and T. montanum L. in xerothermic localities in central and southern Europe (Triberti 1985). The biology of the two species is also similar. No other gracillariid is known to mine in Teucrium (De Prins and De Prins 2018).

In addition, the pupal exuviae of S. botanica (Figs 10, 11) match the description and illustrations of the pupa of Sabulopteryx limosella in the key to Gracillariidae pupae by Patočka and Turčáni (2005: 75–76). The exuviae readily key out to Aspilapteryx in this key, but since the characters of Aspilapteryx were based only on S. limosella, the name Sabulopteryx should be substituted. Characters of S. botanica that lead in this key to Sabulopteryx are as follows: proboscis long, exceeding prothoracic femora; head without projection and rounded in lateral view; pronotum not disconnected on dorsomeson; frontal setae absent; A7 without longitudinal furrows ventrally; A10 with spine-like tubercles. From the description and illustrations in Pereira et al. (2019), the pupa of Vallissiana universitaria Pereira & Arévolo, 2019 shares most of these characters with Sabulopteryx.

Sabulopteryx botanica differs strongly in some characters from its Palaearctic congeners. Neither of the other species has two large spine-like processes on the male valva (Fig. 7); in S. limosella and S. inquinata the single process is on or near the valval costa (Triberti 1985: plate VI B, D); S. botanica has the processes on the valval dorsum. The placement of the ostium in the female genitalia in S. botanica (in the intersegmental membrane between S7 and S8, Fig. 9) is also atypical of Sabulopteryx: in the other species it is at the caudal edge of S7 (Triberti 1985). The invaginations of the intersegmental membrane either side of the ostium (Fig. 9) are apparently unique to S. botanica. The male of S. botanica has T8 in the form of a T-shaped sclerite (Fig. 6), as in genus Aspilapteryx. Given the morphological and molecular support (see below) indicating a close relationship between S. botanica and the other species of Sabulopteryx, these anomalous characters are tentatively considered autapomorphic.

Our molecular analysis, based as it is on a single gene, in no way supplants the much more substantial analysis by Kawahara et al. (2017), but those authors did not include Sabulopteryx (or Aspilapteryx) in their phylogeny. Our analysis (Fig. 20) provides provisional molecular support for the placement of Aspilapteryx and Sabulopteryx in Gracillariinae as suggested by the studies of Kumata (1995), Huemer et al. (2016) and Kawahara et al. (2017), and for the placement of S. botanica in Sabulopteryx, as indicated above from the morphological comparisons.

Figure 20. 

Bayesian consensus phylogeny reconstructed from the COI gene. Numbers above branches represent posterior probabilities. Branch lengths are drawn proportional to the estimated number of substitutions per site, following the scale bar. The tree is rooted with Anthophila fabriciana (Choreutidae).

Despite being only estimated from a single gene, many of the nodes in the phylogeny received posterior probability support values greater than 0.75. Three of the eight subfamilies recovered as monophyletic by Kawahara et al. (2017) are also recovered here, i.e. Gracillariinae (Caloptilia to Mercantouria, posterior probability 0.75, Fig. 20), Acrocercopinae (Spulerina to Artifodina, posterior probability 0.53, Fig. 20), and Parornichinae (Parornix to Callisto, posterior probability 0.68, Fig. 20). Only a single representative each of Marmarinae (Marmara serotinella Busck, 1915), Phyllocnistinae (Phyllocnistis ramulicola Langmaid & Corley, 2007) and Oecophyllembiinae (Eumetriochroa hederae Kumata, 1998) was included. Lithocolletinae (Macrosaccus, Cameraria, Phyllonorycter) were recovered as paraphyletic with respect to Marmarinae, while Ornixolinae (Parectopa, Chileoptila, Micrurapteryx) appeared paraphyletic with respect to Oecophyllembiinae + Phyllocnistinae.

The inconsistencies in our cladogram with respect to Kawahara et al. (2017) are to be expected from a single-gene tree, and suggest the limitations of the current analysis with respect to deeper nodes of the phylogeny. Similarly, there are some inconsistencies with the tree recovered by Huemer et al. (2016), but again these are to be expected and do not undermine the evidence for a close relationship between Sabulopteryx botanica and S. limosella + S. inquinata based on COI, morphology and biology.

The molecular phylogeny supports the treatment of Aspilapteryx and Sabulopteryx as separate genera (Fig. 20), as proposed by Pereira et al. (2019). The two clades are supported as monophyletic with posterior probabilities of 1 and 0.96 respectively. Though the two clades are separated by two nodes that are only weakly supported (0.51 and 0.65 posterior probabilities, Fig. 20), we consider the evidence from the two independent analyses coupled with the differences in morphology and biology outlined by Triberti (1985) and Pereira et al. (2019) as sufficient to warrant full genus status for Sabulopteryx.

Lees et al. (2011) nicely demonstrated the importance of herbarium specimens as a source of information on insect-plant interactions. They searched herbaria for preserved mines of the horse-chestnut leaf miner Cameraria ohridella Deschka & Dimić, 1986 (Gracillariidae: Lithocolletinae), which has recently become invasive throughout Europe, in spite of remaining undetected by entomologists until 1984. From these pressed mines and the larval remains they contained, they were able to document the historical presence of this species in the native range of its host (Aesculus hippocastanum L.) back to 1879, as well as revealing past outbreaks of the moth and novel haplotypes.

Similarly, study of herbarium material (e.g., Fig. 1), in addition to alerting entomologists to the existence of Sabulopteryx botanica, has produced many historical records of the moth. It has greatly helped our knowledge of the distribution and also provided a parasitoid record (see above). The mines are not difficult to find on herbarium sheets, though sometimes magnification is required to scan for the earliest stages. Of 159 herbarium sheets examined in Auckland and Lincoln, 32 (20%) had at least one leaf-mine of S. botanica. The oldest specimen so far found was a single early mine in a leaf from the Cheeseman collection in the Auckland Museum (AK7584): this was collected at Foxhill near Wakefield NN in January 1882. The plant specimen has been annotated appropriately in the Auckland Museum database and the mined leaf is now arrowed on the sheet (E. Cameron, pers. comm.). These old records of the moth also help to confirm that it is an endemic species on its natural host plant and not a recent adventive that has switched to T. parvifolium from an introduced Teucrium species. To check this assumption further, RJBH examined all New Zealand specimens of introduced species of Teucrium (including cultivated species) in the Allan Herbarium in June 2018, and found no evidence of any mines or cocoons. The following species were examined: Teucrium betonicum L’Hér., T. chamaedrys, T. flavum L., T. fruticans L., T. hircanicum L., T. polium L., T. pseudochamaepitys L. and T. scorodonia L. Of these, probably only Teucrium hircanicum and T. scorodonia are established in the wild in New Zealand (NZPCN 2019).