Hoplodactylus duvaucelii ( Duméril & Bibron, 1836 )

Scarsbrook, Lachie, Walton, Kerry, Rawlence, Nicolas J. & Hitchmough, Rodney A., 2023, Revision of the New Zealand gecko genus Hoplodactylus, with the description of a new species, Zootaxa 5228 (3), pp. 267-291 : 272-273

Hoplodactylus duvaucelii ( Duméril & Bibron, 1836)

Figures 2B , 3F–H , 4D–F ; Supplementary Figures 4 , 5D–F

Platydactyle [sic] duvaucelii Duméril & Bibron 1836: 312 ; Duméril & Duméril 1851: 35; Duméril et al. 1854: 248.

Platydactylus duvaucelii Lichtenstein & von Martens 1856: 4 ; Bavay 1869: 6; Boulenger 1883: 126; Chrapliwy et al. 1961: 7.

Hoplodactylus duvaucelii .– Fitzinger 1843: 100; Boulenger 1885: 172; Boulenger 1890: 100; Smith 1933a: 13; Smith 1933b: 377; Stephenson 1948: 339, pl. 63; Hard 1954: 144; Stephenson & Stephenson 1955: 341, figs. 1b, c, 2b, c, 3b, 6a, d, e; McCann 1956a: 46 (in part); Stephenson 1960: 280; Holder 1960: 302 (in part); Myers 1961: 171; Kluge 1967a: 25 (in part); Kluge 1967b: 1013 (in part); Gundy & Wurst 1976: 115; Bauer 1986: 9, figs. 1, 20, 33, 67–69 (in part); Bauer & Russell 1986: 141; Bauer 1987: 593; Worthy 1987a: 219; Worthy 1987b: 416 (in part); Bauer 1990: 108 (in part); Ainsworth et al. 1991: 347; McFadden & Towns 1991: 5; Towns 1991: 125; Worthy 1991: 330; Case et al. 1992: 95; Parrish & Pierce 1993: 57; Bauer & Henle 1994: 139 (in part); Cree 1994: 352 (in part); Daugherty et al. 1994: 318 (in part); Towns 1994: 459; Towns & Daugherty 1994: 327 (in part); Christmas 1995: 4; Towns 1995: 290; Towns et al. 1995: 10; Eifler 1996: 2; Hitchmough 1997: 1 (in part); Towns et al. 1997: 110; Bauer 1998: 43 (in part); Conning & Miller 1999: 32; Atkinson & Towns 2001: 104; Towns et al. 2001: 4 (in part); Towns 2002: 331; Hay et al. 2003: 16 (in part); Parrish & Gill 2003: 209; Todd 2003: 17 (in part); Holmes 2004: 4 (in part); Towns & Atkinson 2004: 11; Hoare 2006: 4, fig. 1; Hoare & Hare 2006: 161; Neilson et al. 2006: 354; Werner & Seifan 2006: 1488; Barry et al. 2007: 260; Gardner-Gee et al. 2007: 32; Hoare et al. 2007: 511, fig. 1; Ji et al. 2007: 264; van Winkel et al. 2007: 271; Moore et al. 2008: 457; Todd 2008: 105; van Winkel 2008: 15, pls. 1.1, 2.1–2.2, 3.1–3.2, 4.1–4.3, 5.1, 7.1; Agnew 2009: 19, figs. 24–25; Bell 2009: 417, fig. 2; Lee et al. 2009: 834; Towns et al. 2009: 14; Barry 2010: 2; Barry et al. 2010: 235; Bellingham et al. 2010: 135; Gardner-Gee & Beggs 2010: 296, fig. 3; Middleton et al. 2010: 249; Russell et al. 2010: 170, fig. 9; van Winkel et al. 2010: 113, fig. 1; Barry et al. 2011: 199, fig. 1; Hare 2011: 274; Murphy & Thompson 2011: 578; Nielsen et al. 2011: 17; fig. 7 (in part); Shea et al. 2011: 5; Wong et al. 2011: 331; Dhami et al. 2012: 538; Moir et al. 2012: 203; Remeš et al. 2012: 55; Thomlinson 2012: 8; van Winkel & Ji 2012: 203; Armstrong & Ewen 2013: 290; Baling et al. 2013a: 250; Baling et al. 2013b: 275; Berner et al. 2013: 466; Galbraith & Cooper 2013: 259; Hitchmough et al. 2013: 10 (in part); Jones et al. 2013: 41; Parker 2013: 286; Barry et al. 2014: 396; Bell 2014: 8 (in part); Daza et al. 2014: 443, figs. 11g –i; Jarvie & Monks 2014: 211; Miller et al. 2014: 1048; Monks et al. 2014: 169; Nichols 2014: 10; Romijn et al. 2014: 111 (in part); van Winkel & Weihong 2014: 13, fig. 2; Dumont 2015: 8; Evans et al. 2015: 263; Gibson et al. 2015: 891; Heath & Whitaker 2015: 751 (in part); Holdom 2015: 8, figs. 1.1, 2.6, 2.7, 3.1, 5.4; Le Souëf et al. 2015: 340, figs. 1–3; Mockett 2015: 73 (in part); Brown et al. 2016: 38; Chapple 2016a: 371; Chapple 2016b: 4 (in part); Chapple & Hitchmough 2016: 116 (in part); Cree & Hare 2016: 174 (in part); Gartrell 2016: 213; Gardner et al. 2016: 929; Glenday 2016: 3, pls.1.0, 2.0, 2.1a–2.1c, 2.2a, 2.2b, 3.0–3.3, 3.4a–3.4c, 4.1, 5.1–5.6, 6.0; Gollin 2016: 54; Hare and Cree 2016: 246 (in part); Hare et al. 2016: 140, fig. 6.3 (in part); Hitchmough et al. 2016a: 9 (in part); Hitchmough et al. 2016b: 89 (in part); Lettink & Hare 2016: 17; Morgan-Richards et al. 2016: 77, fig. 2 (in part); Nelson et al. 2016: 333; Paluh 2016: 30, fig. 11i; Plein et al. 2016: 1187; Romijn & Hartley 2016: 196 (in part); Scott 2016: 16; Shea 2016: 18; Towns et al. 2016a: 242; Towns et al. 2016c: 123; Worthy 2016: 71 (in part); Harker et al. 2017: 306, fig. 1; Lozito & Tuan 2017: 148, fig. 2 (in part); Mockett 2017: 42; Vasconcelos et al. 2017: 8; Busbridge & Stewart 2018: 192; Paluh & Bauer 2018: 698; Sullivan 2018: 74; van Winkel et al. 2018: 114, pls. 35, 40, 46, 127 (in part); Skipwith et al. 2019: 10 (in part); While et al. 2019: 332; Woolley et al. 2019: 2; Andruzzi et al. 2020: 2; Glynne et al. 2020: 804 (in part); Woolley 2020: 15; Gollin et al. 2021: 7; Elangovan et al. 2021: 1; Scarsbrook 2021: 19, figs. 1.5, 2.1, 3.1, 4.1 (in part); Scarsbrook et al. 2021: 2 (in part); Scarsbrook et al. 2022: 3, fig. 1 (in part).

Naultinus pacificus .– Gray 1843: 203 (in part); Blyth 1859: 279; Buller 1870: 7 (not Gray, 1842).

Pentadactylus duvaucelii .– Gray 1845: 160; Günther 1864: 118.

Hoplodactylus granulatus .– Lucas & Frost 1897: 273 (in part, not Gray, 1845).

Hoplodactylus duvancellii [sic].– Schaefer 1902: 35.

Haplodactylus duvaucelii .– Womersley 1941: 328.

Rhacodactylus trachyrhynchus .– Guibé 1954: 16 (not Bocage, 1873).

Hoplodactylus duvaucellii [sic].– Hard 1954: 144; Towns 1971: 91.

Hoplodactylus duvauceli [sic].– McCann 1955: 39, fig. 3, pl. 4 (in part); McCann 1956a: 46 [in part]; McCann 1956b: 15 (in part); Kinsky & Sibson 1959: 137; Atkinson 1964: 399; Atkinson 1965: 3; Sharell 1966: 49, pls. 28–31 (in part); Thoresen 1967: 197; Atkinson 1968: 287; Merton & Atkinson 1968: 107; Whitaker 1968: 623; Hardy 1972: 165; Richards 1973: 228; Towns & Hayward 1973: 94; Whitaker 1973: 122; Towns 1974: 35; Hicks et al. 1975: 211; Bell 1976: 318; McCallum 1981a: 153; McCallum 1981b: 55; Ogle 1981: 192; Porter 1981: 10; McCallum & Harker 1982: 22; Harper 1983: 307; Bau- er 1985: 90 (in part); Newman & Towns 1985: 279; Whitaker 1987: 315; Holdaway 1989: 12. (unjustified emendation).

Naultinus duvaucelii .– Chrapliwy et al. 1961: 6 (in part).

Hoplodactylus diwancelii [sic].– Jullien & Renous-Lécuru 1973: 14.

Hoplodactylus taranganus Steindachner in Bauer 1987: 594 (nomen nudum).

Woodworthia duvaucelii .– Jewell 2008: 50 (in part).

Type material.— Lectotype MNHN-RA 5977 and paralectotypes ( MNHN-RA 6680 , MNHN-RA 6681 , RMNH 2722 ), “Bengal” (in error, = New Zealand). Type locality (by subsequent unintended designation of Bauer, 1990, see below): Taranga Island , Hen and Chicken Islands.

Material examined.— The lectotype from images only; not the paralectotypes. “Cape Maria van Dieman” ( OMVT925 , OMVT939 , OMVT940 ). Poor Knights Islands: Tawhiti Rahi Island (RE.003534, RE.003535); Aorangi Island (RE.001751, RE.006490, CD1032). Bream Island (RE.002674). Hen and Chicken Islands : Muriwhenua Island (RE.003491); Coppermine Island (FT630); Taranga Island (RE.006492, FT576, FT580). Great Barrier Island (RE.003011, LH3223). Mercury Islands : Stanley Island (RE.003256); Double Island (RE.003160); Middle Island (FT175, FT176); Green Island (FT177, FT178); Korapuki Island (RE.003157, FT179). Alderman Islands : Hernia Island (RE.006671, FT560); Raumahuanui Island (FT566); Hongiora Island (FT567); Middle Chain Island (FT562); Raumahuaiti (FT564). Waikato (mainland): Maungatautari (RE.007381); precise locality unknown, historic ( LH166 ). Waitomo (mainland): Holocene fossil (AU7700.2; WO333). Wairarapa (mainland): Mataikona River , Holocene fossil (S.46528.2); Ruakokopotuna , Holocene fossil (S.47439) .

Distribution.— New Zealand: formerly throughout the North Island (Holocene); presently restricted to islands off the north-eastern North Island and some mainland predator-free sanctuaries ( Fig. 1 ).

Remarks.— The type series for H. duvaucelii were labelled simply “Bengal, Duvaucel”, with the Indian provenance clearly an error ( Smith 1933a, b; Stephenson 1948; Bauer 1987, 1990). Smith (1933b) subsequently designated New Zealand as the type locality (ICZN Art. 76A), then Bauer (1990) unintentionally designated the Hen and Chicken Islands as the revised type locality, misciting Smith (1933b). The lectotype is clearly from the North Island or a nearshore island off same (pers. obs.; Bauer 1987; Supp. Fig. 4 ), and the Hen and Chicken Islands are a plausible source population for the type series. This is both the location mentioned in Smith (1933b) and, fortuitously, the inferred type locality (i.e., Taranga [Hen] Island) of a nomen nudum based on a series of specimens at NHW (“ Hoplodactylus taranganus Steind. ”; see: Bauer 1987).

The provenance of the “Cape Maria van Dieman” (CMVD) specimens is unclear. That name has variously, informally been applied to Motuopao Island (off CMVD), or even much of the Aupouri Peninsula/Te Hiku o te Ika (on which CMVD is located) in some contexts (especially on historical specimen labels), rather than strictly the tiny headland of CMVD itself. Additionally, many museum specimens dating from around the turn of the 20 th century originated from lighthouse keepers, who traded with visiting vessels and naturalists as well as their local communities, in addition to collecting samples themselves (KW pers. obs.). It is possible this provenance refers to the samples coming to OM from the keepers of the Motuopao Island or CMVD lighthouses, and not necessarily the actual origin of the samples themselves (Kane Fleury pers. comm. 2021). However, these specimens are genetically distinct from all other sampled North Island populations ( Scarsbrook et al. 2022), and therefore, a Far North origin seems probable (if not from CMVD itself). It should also be noted that the northernmost extant island populations of H. duvaucelii (Stephenson’s Island and Cavalli Islands) have not been sampled genetically.

Although the origin of the enigmatic Maungatautari specimen (RE.007381; Morgan-Richards et al. 2016) remains unknown, we consider it more likely to be an escapee from a captive population rather than from a relictual, otherwise unknown natural mainland population. Genetically the sample is unique but close to those from islands in the Bay of Plenty ( Scarsbrook et al. 2022). No other verified mainland sightings of living H. duvaucelii have been reported in at least the last 60 years.

Considerable genetic ( Scarsbrook et al. 2022) and morphological ( Scarsbrook et al. 2021) variation occurs among the numerous extant populations of H. duvaucelii . Those from Great Barrier Island (now extremely rare, possibly extinct) and the Poor Knights Islands are especially divergent and undoubtably warrant recognition as distinct management units to maximize preservation of remaining diversity within this species. Intensive and urgent monitoring surveys should therefore be carried out on Great Barrier Island to establish whether a low-density population of H. duvaucelii persists (last reported capture in 2011; Morgan-Richards et al. 2016). Specimens from the Poor Knights Islands, where this species occurs in high densities ( McCallum 1981a), are morphologically highly variable relative to other extant populations, and notably include a high proportion of relatively large, pale and lightly patterned specimens. Historical and often undocumented translocations of this species, or museum labelling errors, may have slightly confounded reported phylogenetic signals ( Scarsbrook et al. 2022) as well as interpretations of morphological characters. Future translocation events and management of captive populations should be informed by these results.

Suggested IUCN Red List status of H. duvaucelii is ‘ Critically Endangered A 1 (a, b, c, e)’: population reduction of>90% observed, estimated, inferred, or suspected in the past where the causes of the reduction are clearly reversible AND understood AND have ceased. This is based on range contractions (following the arrival of humans in New Zealand), from widespread and abundant on the mainland (i.e., Northland to Wairarapa ; Fig. 1 ), to presently restricted to small islands and predator-free mainland sanctuaries (with the completion of mainland extirpation probably having occurred during the 20 th century, within the last three generations for this very long-lived species) .

Figures

FIGURE 1. Maps of New Zealand showing distribution of Hoplodactylus duvaucelii (Duméril & Bibron 1836) (left) and H. tohu n. sp. (right), with inter- and intraspecific phylogenetic distance highlighted in a simplified mitochondrial genome phylogeny (bottom; adapted from Scarsbrook et al. 2022). Extant relict (pink) and translocated (purple) populations, historic (yellow) and Holocene subfossil records (blue) are indicated (Supp. Tables 1–2), with triangles denoting localities included in genetic analyses (see phylogeny). Abbreviations for referenced localities are as follows: Cape Maria van Dieman (CM), Poor Knights Islands (PK), Hen and Chickens Islands (HC), Great Barrier Island (GB), Maungatautari Ecosanctuary (ME) and Mana Island (MI).

FIGURE 2. Hoplodactylus (Fitzinger, 1843) sp.—dorsal and ventral body views (left-right). A, Hoplodactylus tohu n. sp., holotype, Middle Trios Island, Marlborough Sounds, RE.000503. B, Hoplodactylus duvaucelii Duméril & Bibron, 1836, topotype, Muriwhenua Island, Hen and Chickens Islands, RE.003491. Automontage images generated by Jean-Claude Stahl (NMNZ). Scale bar = 20 mm.

FIGURE 3. Details of: A-D, Hoplodactylus tohu n. sp., holotype (RE.000503); and E-H, Hoplodactylus duvaucelii Duméril & Bibron, 1836, topotype (RE.003491): A, E, right-lateral view of head; B, F, oblique right-lateroventral view of head, with diagnostic characters (a) pronounced brillar fold, and (b) abrupt size decrease at the 5th infralabial scale; C, G, ventral view of the right manus showing subdigital lamellae, and highlighting the less emergent claw on the first digit in H. duvaucelii; D, H, ventral view of the right pes showing subdigital lamellae. Automontage images generated by Jean-Claude Stahl (NMNZ). Scale bar = 10 mm.

FIGURE 4. Living individuals of Hoplodactylus species. A–C, Hoplodactylus tohu n. sp.; D–F, H. duvaucelii (Duméril & Bibron 1836). A, Brothers Islands (Cook Strait); B–C, Sentinel Rock (Marlborough Sounds). D–F, Korapuki Island (Mercury Islands). The first and second rows show sub-adult and adult representatives of each species respectively. Photographs: Nicholas Harker (Auckland, New Zealand).