Lichen notes from the Kermadec Islands. II. Ramalina

Abstract

Eleven species of Ramalina ( R. australiensis, R. canariensis, R. celastri, R. exiguella, R. geniculata, R. leiodea, R. luciae, R. meridionalis, R. microspora, R. pacifica and R. peruviana) are accepted for the Kermadec Islands. The northern Kermadec Islands had the greatest diversity of species (eleven species) while the southern Kermadec Islands had collectively five species. These were distributed as follows (islands arranged from north to south): northern Kermadec Islands (the Herald Islets (six species), the Meyer Islands group (two species), Raoul Island (nine species)); southern Kermadec Islands (Macauley Island (six species), Curtis Island (one species), Cheeseman Island (one species), L’Esperance (two species)). Ramalina leiodea is a new addition to the mycobiota of the New Zealand Botanical Region and R. canariensis and R. meridionalis are new records for the Kermadec Islands, and a significant easterly extension for R. meridionalis.

Full text

Bulletin of the Auckland Museum 20: http://www.aucklandmuseum.com/research/pub/bulletin/20/5
INTRODUCTION
The rst apparent collections of Ramalina from the
Kermadec Islands (Fig. 1) were made by W.R.B. Oliver in
1908. Those collections, along with all of Oliver’s other
lichen specimens, were identied by A.D. Cotton at Kew
and the results were then published by Oliver (1912). In
that paper Oliver reported two species, Ramalina fastigiata
Ach. and R. farinacea Ach. from “Sunday Island” (Raoul
Island). However, subsequent examination of Oliver’s
specimens in New Zealand herbaria, indicate that they
are referable to R. microspora Kremp. (WELT L04263),
R. pacica Asahina (AK 202677, WELT L04265) and R.
peruviana Ach. (AK 33480).
Lichen collections were also made by R.C. Cooper
from Raoul Island in 1956, and they, along with Oliver’s
1908 collections and the more extensive ones made
by W.R. Sykes, were examined by T.W. Rawson, who
did all the lichen determinations listed for the islands
published by Sykes (1977) in his Flora of the Kermadec
Islands (see de Lange & Galloway 2015). Sykes (1977)
listed six species for the northern Kermadec Islands and
one for the southern Kermadec Islands. The species
listed by Sykes (1977) were: Ramalina allanii B. de
Lesd. (= R. australiensis Nyl.), R. farinacea (specimens
are R. pacica (WELT L4265) or R. peruviana (AK
304651)), R. fastigiata (specimen (WELT L4263)
is R. microspora), R. intermediella Vain. (specimen
(CHR 385510) is R. luciae Molho, Bodo, W.L.Culb. et
C.F.Culb.), R. microspora and R. pacica.
Blanchon et al. (1996) listed six species for the
Kermadec Islands (R. australiensis, R. exiguella Stirt.,
R. geniculata Hook.f. et Taylor, R. luciae, R. pacica and
R. peruviana), based on the original W.R.B. Oliver, R.C.
Cooper and W.R. Sykes collections noted above and also
unaccessioned Raoul Island material collected by C.J.
West (from a ten-week trip she made there in 1993).
Elix and McCarthy (1998) subsequently listed eight
species for the Kermadec Islands, largely following
Blanchon et al. (1996), recognising R. australiensis,
R.exiguella, R. luciae, R. pacica and R. peruviana but
omitting R. geniculata. They also accepted without
comment R. farinacea, R. fastigiata and R. microspora
from Sykes (1977).
Bannister and Blanchon (2003), examining W.R.B.
Oliver, R.C. Cooper and W.R. Sykes’ Kermadec Islands
Ramalina material, and also unaccessioned material
collected by B.W. Hayward and A.E. Wright from
Raoul Island in 1994, accepted eight species: Ramalina
australiensis, R. exiguella, R. geniculata, R. luciae, R.
microspora (as R. cf. microspora), R. pacica, and R.
peruviana for the islands, and they added R. celastri
(Spreng.) Krog et Swinscow. They identied a specimen
(AK 202677) collected by W.R.B. Oliver in 1908 and
Abstract
Eleven species of Ramalina (R. australiensis, R. canariensis, R. celastri, R. exiguella, R.
geniculata, R. leiodea, R. luciae, R. meridionalis, R. microspora, R. pacica and R. peruviana)
are accepted for the Kermadec Islands. The northern Kermadec Islands had the greatest diversity
of species (eleven species) while the southern Kermadec Islands had collectively ve species.
These were distributed as follows (islands arranged from north to south): northern Kermadec
Islands (the Herald Islets (six species), the Meyer Islands group (two species), Raoul Island (nine
species)); southern Kermadec Islands (Macauley Island (six species), Curtis Island (one species),
Cheeseman Island (one species), L’Esperance (two species)). Ramalina leiodea is a new addition
to the mycobiota of the New Zealand Botanical Region and R. canariensis and R. meridionalis
are new records for the Kermadec Islands, and a signicant easterly extension for R. meridionalis.
Keywords
Kermadec Islands; Ramalinaceae; Ramalina; R. australiensis; R. canariensis; R. celastri;
R. exiguella; R. geniculata; R. leiodea; R. luciae; R. meridionalis; R. microspora; R. pacica;
R. peruviana; biogeography; New Zealand botanical region.
Lichen notes from the Kermadec Islands.
II. Ramalina
Peter J. de Lange Department of Conservation
Dan J. Blanchon Unitec Institute of Technology
171–181

172
originally determined as R. farinacea as R. pacica, and
further noted that another specimen collected by Oliver
and labelled as R. fastigiata, contained divaricatic acid
and could belong to R. cf. microspora (Bannister &
Blanchon 2003).
The most recent lichen ora treatment for New Zealand
(Galloway 2007) recognised seven species of Ramalina
for the Kermadec Islands, listing R. australiensis, R.
celastri, R. exiguella, R. luciae, R. pacica and R.
peruviana from Raoul Island, and R. geniculata from the
Kermadec Islands in general. Notably, Galloway (2007)
did not accept Ramalina microspora for the islands,
noting (Pp. 1506) that “a fertile, non-sorediate taxon (cf.
R. microspora Kremp.) is also known from coastal lava
outcrops in scrub from Raoul Island in the Kermadec
Islands (Bannister & Blanchon 2003). But its exact status
is still in doubt and it is not included here”. During May
2011, the senior author visited the Kermadec Islands and
collected specimens of Ramalina from Raoul Island, the
Meyer Islands and Herald Islets, Macauley Island, Curtis
Island, Cheeseman Island and L’Esperance Rock (Fig. 1).
From those collections Blanchon et al. (2012) have now
conrmed the presence of Ramalina microspora on the
Kermadec Islands.
Eleven species are now known from the Kermadec
Islands. Three species, R. canariensis Steiner, R. leiodea
(Nyl.) Nyl. and R. meridionalis Blanchon et Bannister,
are new to the Islands. Ramalina leiodea is also a new
addition to the New Zealand Botanical Region (sensu
Allan 1961, de Lange & Rolfe 2010) mycobiota.
Because the other Ramalina collections signicantly
expand the known range of the genus on the Kermadec
Islands, we provide here a full treatment and key to the
genus Ramalina for the islands.
Peter J. de Lange & Dan J. Blanchon
Figure 1. The Kermadec Islands. A. Location of the Kermadec Islands in relation to the continental islands of New Zealand.
B. Spatial relationship between the northern and southern islands of the Kermadec Islands. C. Raoul Island, the Meyer and
Herald Islets – collectively the northern Kermadec Islands. Map by Jeremy R. Rolfe

173
ARTIFICIAL KEY TO SPECIES OF RAMALINA ON THE KERMADEC ISLANDS
THE SPECIES
Ramalina australiensis Nyl.
Description (Fig. 2) Thallus corticolous or saxicolous,
yellow-green, green to grey-green, erect to pendulous,
30–150 mm long, rarely to 250 mm; branching sparse
in basal region to dense and irregular towards the apices
where small branchlets emerge at right angles to the
primary branch; branch width 0.1–1.5(–2.0) mm, primary
branches subterete and ne or attened and coarse,
ne secondary branches often occur at right angles to
main branch, apices curled or hooked; surface matt,
longitudinally grooved near base; linear pseudocyphellae
often present; holdfast delimited; soralia absent.
Apothecia uncommon, sessile, lateral, +/– basal, plane
to convex, disc 0.5–1.0(–1.3) mm diam., margin entire,
pale pink-orange; ascospores ellipsoid, straight, 12–16 ×
4.0–6.0 µm. Chemistry: Usnic acid only. Medulla K–
Description based on Blanchon et al. (1996).
Northern Kermadec Islands: Raoul Island: Five
specimens seen: Hutchinson Bluff, W.R. Sykes 412/K,
5 Dec 1966, CHR 352164; East of Coral Bay, B.W.
Hayward KE36, 26 Nov 1994, AK 224388; Boat Cove,
A.E. Wright 12486, 26 Nov 1994, AK 224392; Green
Lake, B.W. Hayward KE5, 21 Nov 1994, AK 224398;
Near bottom of Sunshine Valley, B.W. Hayward KE29,
24 Nov 1994, AK 224402. Herald Islets: One specimen
seen: Dayrell Island, P.J. de Lange K1279, 18 May 2011,
UNITEC 6953. Southern Kermadec Islands: Macauley
Island: One specimen seen: “Homalanthus Bush”, P. J.
de Lange K1200, 21 May 2011, UNITEC 5912.
Comments Ramalina australiensis has been collected
from Raoul and Dayrell Islands in the northern Kermadec
Islands, and from Macauley Island in the Southern
Kermadec Islands. In the Northern Kermadec Islands,
specimens were collected from Kermadec pohutukawa
(Metrosideros kermadecensis) and from Macauley Island,
on Kermadec popular (Homolanthus populifolius) .
Figure 2. Ramalina australiensis, Raoul Island, Green Lake
(B.W. Hayward KE5, AK 224398). Scale Bar 20 mm.
Photo: Dan Blanchon
Lichen notes from the Kermadec Islands. II. Ramalina

174 Peter J. de Lange & Dan J. Blanchon
Corticolous specimens tend to have long, ne branches,
and can be confused with ne specimens of R. peruviana
or R. geniculata. However, R. peruviana is sorediate and
has a chemistry of the sekikaic acid aggregate, whereas R.
geniculata is hollow and perforate. Saxicolous specimens
can resemble R. meridionalis but that species branches
mainly from the base and has norstictic acid in its medulla
(K+ red), whereas R. australiensis has denser branching
of ne side branches towards the apices and is K–.
Ramalina canariensis Steiner
Description (Fig. 3) Thallus corticolous, pale-green to
yellow-green, erect, 8–35mm long; branching palmate
or irregular; branch width 1.0–3.5 mm, branches at,
apices blunt, usually split; surface matt, smooth to
rugose; holdfast delimited; soralia marginal and apical,
often developing with the separation of upper and lower
surfaces. Apothecia not seen in Kermadec or New
Zealand material. Chemistry: Usnic acid and divaricatic
acid. Medulla K–.
Description based on Blanchon et al. (1996).
Northern Kermadec Islands: Raoul Island: One
specimen seen: Sunshine Valley, P. J. de Lange K1206,
16 May 2011, UNITEC 5967.
Comments Ramalina canariensis is a mainly maritime
species, with Stevens (1987) recording it from parts of
Australia with a Mediterranean type of climate, and also
from the Mediterranean region, Canary Islands, South
Africa and South America. In New Zealand the species
has been referred to as having a subtropical maritime
distribution (Blanchon et al. 1996), but Bannister et al.
(2004) noted that specimens had been found in inland
sites in New Zealand at 260 m in altitude. In New
Zealand the species is not commonly seen, resulting
in its current listing as ‘Naturally Uncommon’ by the
New Zealand Lichen Threat Listing Panel (de Lange
et al. 2012).
Ramalina celastri (Spreng.) Krog et Swinscow
Description (Fig. 4) Thallus corticolous or saxicolous,
pale green to grey-green, occasionally blue-grey, erect
or pendulous, 10–120(–150) mm long; branching
trichotomous, subdichotomous or irregular, small
branches at right angles to the main branch sometimes
present; branch width (1–)3–10 mm, occasionally to
20 mm, branches range from narrow and canaliculate
with ne apices to broad and at with blunt or tattered
tips; surface matt, smooth to ridged or rugose, thin to
coarse and thick, often torn or fenestrate in older thalli,
sometimes producing small lobules from torn areas;
pseudocyphellae common, short linear to punctiform;
holdfast delimited; soralia absent. Apothecia numerous,
marginal on narrow branches, marginal and laminal on
broader branches; disc 0.3–2.0(–4.0) mm in diam., plane
to convex; margin narrow, entire, smooth or incised,
often occluded with age; ascospores oval, ellipsoid,
straight, 9.5–15.5(–17.5) × 4.0–6.0 µm. Chemistry:
Usnic acid and two unknowns (trace). Medulla K–.
Description based on Blanchon et al. (1996).
Northern Kermadec Islands: Raoul Island: Two
specimens seen: Green Lake, B.W. Hayward KE5, 21 Nov
1994, AK 224397; Denham Bay Track, Orange Grove
Track, P.J. de Lange K749, 18 May 2011, AK 330764.
Herald Islets: One specimen seen: Dayrell Island, P.J. de
Lange K1278, 18 May 2011, UNITEC 6955.
Comments Ramalina celastri, a widespread,
pan-subtropical species (Stevens 1987, Blanchon et al.
1996), has been recorded only from Raoul and Dayrell
Islands. This photophilous species is evidently very
uncommon on Raoul Island. Its absence from historical
lichen collections made from Raoul Island, at a time
when the island’s forest was much degraded through goat
browse, and as a result frequented by photophilous lichen
species such as Ramalina pacica and Crocodia aurata
(Ach.) Link (de Lange & Galloway 2015), suggests
that R. celastri could even be a recent arrival to Raoul.
Figure 3. Ramalina canariensis, Raoul Island, Sunshine
Valley (P. J. de Lange K1206, UNITEC 5967). Scale Bar
20 mm. Photo: Dan Blanchon
Figure 4. Ramalina celastri, Raoul Island, Green Lake (B.W.
Hayward KE5, AK 224397). Scale Bar 20 mm.
Photo: Dan Blanchon

175
On Raoul Island the species appears to be exclusively
corticolous, although in New Zealand it is routinely
either saxicolous or corticolous (Blanchon et al. 1996,
Hayward et al. 2014). The only species R. celastri is
likely to be confused with is R. leiodea but that species
is usually canaliculate, has concave or plane apothecia,
and has a boninic acid chemistry, whereas R. celastri has
plane to convex apothecia and no medullary chemistry.
Ramalina exiguella Stirton
Description (Fig. 5) Thallus corticolous, grey-green
to yellow-green, erect, rigid, 8–15(–20) mm long;
branching mainly basal but with short, lateral branches
appearing in some specimens; branch width 0.2–0.6 mm,
branches subterete to terete, narrow, apices ne; surface
shiny or matt, regularly grooved to cause a string-like
appearance; pseudocyphellae common, linear; holdfast
delimited; soralia absent. Apothecia common, few to
many, marginal and subterminal causing branch tips
to become geniculate; disc 0.3–1.7 mm diam., plane
to convex; margin entire; ascospores ellipsoid, bent
or rarely straight, occasionally reniform, 11.5–15.5 ×
5.0–7.0 µm. Chemistry: Usnic acid only. Medulla K–
Description based on Blanchon et al. (1996).
Northern Kermadec Islands: Raoul Island: Five
specimens seen: Green Lake, B.W. Hayward KE5, 21
Nov 1994, AK 224387; Green Lake, B.W. Hayward
KE5, 21 Nov 1994, AK 224396; Lava Point, A.E. Wright
12446, 23 Nov 1994, AK 224395; Green Crater, C.J.
West s.n., 1 Jul 1993, WELT L04403; Sunshine Coast,
Near Boat Cove, 21 May 2012, P.J. de Lange K1160, AK
330765; Herald Islets: One specimen seen: Dayrell
Island, P.J. de Lange K553, 18 May 2011, AK 330754.
Comments Ramalina exiguella, a typically corticolous,
photophilous species from East Africa, Australia and
New Zealand (Blanchon et al. 1996), is known in the
Kermadec Islands only from the northern Kermadec
Islands, where it has only been recorded from Raoul
and Dayrell Islands. With one exception, the saxicolous
gathering of P.J. de Lange K1160, the Kermadec
collections were all corticolous, being collected from the
exposed twigs and branchlets of Kermadec pōhutukawa.
On the Kermadec Islands this species is likely to be
confused with only R. geniculata but it is not hollow and
perforate, and it does not have any medullary chemistry.
Ramalina geniculata Hook.f. et Taylor
Description (Fig. 6) Thallus corticolous and saxicolous,
green to yellow-green, erect to subpendulous, 20–40(–50)
mm high; branching dichotomous to subdichotomous;
branch width 0.5–2.0(–4.0) mm, branches hollow,
perforate, perforations oval; surface matt to shiny,
smooth to rugose; pseudocyphellae rare, pustulate;
holdfast delimited; soralia absent. Apothecia common,
terminal or subterminal, usually geniculate (spurred);
disc 0.5–3.0(–3.5) mm diam., plane or convex; margin
entire but occluded at maturity; ascospores ellipsoid,
straight or slightly curved, (8.5–)9.5–13.5 × 4.0–7.0
µm. Chemistry: Usnic acid, sekikaic acid (major) +/–
divaricatic acid (minor), or divaricatic acid (major) +/–
sekikaic acid. K+ pink.
Description based on Blanchon et al. (1996).
Northern Kermadec Islands: Raoul Island: Nine
specimens seen: Bell’s Terrace, C.J. West s.n., Jul
1993, WELT L04405; Green Lake, B.W. Hayward KE5,
21 Nov 1994, AK 224387; Northeast end of Denham
Bay, B.W. Hayward KE12, 22 Nov 1994, AK 224390;
near bottom of Sunshine Valley, B.W. Hayward KE29,
24 Nov 1994, AK 224401; Mouth of Sunshine Valley,
B.W. Hayward KE30, 24 Nov 1994, AK 224389;
Sunshine Valley, A.E. Wright 12473, 24 Nov 1994, AK
224391; North Terraces, B.W. Hayward KE34, 25 Nov
1994, AK 224399; Lava Point, A.E. Wright 12490,
26 Nov 1994, AK 224393; Bluff near Boat Cove, P.J.
de Lange K547, 16 May 2011, AK 330747. Southern
Kermadec Islands: Macauley Island: Two specimens
seen: Western slopes of Mt Haszard, W.R. Sykes 1054/K,
Figure 5. Ramalina exiguella, Herald Islets, Dayrell Island
(P.J. de Lange K553 AK 330754). Scale Bar 20 mm.
Photo: Dan Blanchon
Figure 6. Ramalina geniculata, Raoul Island, Sunshine
Valley (B.W. Hayward KE29 AK 224401). Scale Bar 20 mm.
Photo: Dan Blanchon
Lichen notes from the Kermadec Islands. II. Ramalina

176 Peter J. de Lange & Dan J. Blanchon
18 Nov 1970, CHR 211875; Mt Haszard, P.J. de Lange
K556, 21 May 2011, AK 330757.
Comments Ramalina geniculata is a corticolous and
saxicolous species also found in New Zealand (Blanchon
et al. 1996). Fine forms such as P.J. de Lange K547 (AK
330747) approach the morphology of R. australiensis,
R. exiguella or R. peruviana, and more robust forms
like those collected from Macauley Island can resemble
R. microspora. The hollow and perforate nature of
R. geniculata easily distinguishes it from those other
species. In the northern Kermadec Islands, R. geniculata
was collected only from Raoul Island, mostly from open,
coastal sites and from boulders lying in rank pasture.
Despite the abundance of this habitat on the nearby
Meyer Islands and the Herald Islets, the species was not
seen there during the 2011 survey. On Macauley Island,
R.geniculata seems to be uncommon, though there it grew
with the supercially similar R. microspora and as such it
might have been confused with that species in the eld.
Ramalina leiodea (Nyl.) Nyl.
Description (Fig. 7) Thallus corticolous, grey-green,
erect or subpendulous, 40–80(–90) mm long;
branching subdichotomous, sparse to moderate;
branch width (0.5–)2.0–4.0 mm, branches compressed,
at or distinctly canaliculate when narrow, apices
attenuate; surface matt, smooth to rugose, short
linear pseudocyphellae sometimes present; holdfast
delimited; soralia absent. Apothecia common, marginal
and subterminal, rarely laminal, small thalli produce
subterminal apothecia subtended by a spur; disc 1–3
mm in diam., concave to plane; margin entire or
incised at maturity; ascospores oval, ellipsoid, straight,
or slightly curved, 9.5–15.5(–17.5) x 4.0–6.0 µm.
Chemistry: Boninic acid (major), 2-0-methylsekikaic
acid (minor), 2,4'-di-0-methylnorsekikaic acid
(minor), 4'-0-methylpaludosic acid (minor),
4,4'-di-0-methylcryptochlorophaic acid (minor), and
usnic acid. Medulla K–.
Description based on Stevens (1987).
Northern Kermadec Islands: Herald Islets: Two
specimens seen: North Chanter, South summit, P.J. de
Lange K620, 16 May 2011, AK 330761; Dayrell Island,
P.J. de Lange K1289, 18 May 2011, UNITEC 6954.
Southern Kermadec Islands: Macauley Island: One
specimen seen: Mt Haszard, western seaward cliffs, P.J.
de Lange K557, 21 May 2011, AK 330758.
Comments Ramalina leiodea is recorded from islands
in the Indian and Pacic Oceans, Australia, New
Caledonia, Norfolk Island and Lord Howe Island
(Stevens 1987). Its presence therefore on the Kermadec
Islands is to be expected, and it is indeed surprising it
has not been recorded from there before. The current
distribution may in part explain this, as neither Dayrell,
North Chanter nor Macauley Island are often visited,
and then rarely so by botanists. Interestingly, Sykes
(1977) collected the saxicolous R. microspora from
Macauley Island and it is tempting to speculate that had
R. leiodea been present then, because it is a conspicuous
species, he would almost certainly have gathered it too.
However, the type of habitat in which it was collected on
Macauley Island, Kermadec ngaio (Myoporum rapense
subsp. kermadecense (Sykes) Chinnock) treeland was
Figure 7. Ramalina leiodea, Macauley Island, Mt Haszard
(seaward clis) (P.J. de Lange K557, AK 330758). Scale
Bar 20 mm. Photo: Dan Blanchon
Figure 8. Ramalina luciae, Raoul Island, road to Ravine 8
(W.R. Sykes 679/K, CHR 385510). Scale Bar 20 mm.
Photo: Dan Blanchon

177
scarce during Syke’s visits due to goat browse. Had it
been present, then it is likely that it probably grew in
cliff refugia on shrubs and trees, virtually inaccessible
to humans. The only species likely to be confused with
R. leiodea is R. celastri but that species lacks boninic
acid and tends to have convex apothecia. An earlier
Raoul Island collection (A. E. Wright 12490, (AK
284975)) could also be R. leiodea; the specimen is very
fragmentary but boninic acid was detected with thin
layer chromatography (J. Bannister pers. comm.).
Ramalina luciae Molho, Bodo, Culb. et C.Culb.
Description (Fig. 8) Thallus corticolous, pale green or
green-yellow, subpendulous to pendulous, 44–120(–190)
mm long; branching dichotomous, elongation between
branching; branch width 0.5–3.0 mm, branches attened
at base and subterete at apices, growth form tangled
due to fusion of branches; cortex matt; pseudocyphellae
present, linear, inconspicuous; soralia marginal and
laminal, round to ellipsoid. Apothecia not seen in New
Zealand material. Chemistry: Usnic acid, sekikaic
acid (major), 4'-O-demethylsekikaic acid (minor),
4'-O-methylnorsekikaic acid (minor) and ramalinolic
acid (trace). Medulla K+ pink.
Description based on Blanchon et al. (1996).
Raoul Island: Five specimens seen: Between
Meteorological Station, Boat Cove and Fishing Rock,
R.C. Cooper 44209, 44289, Jun 1956, AK 310856, AK
310857; Above farm, W.R. Sykes 679/K, 3 Jan 1967,
CHR 385510;, Beside road to Ravine 8, Bell’s Flat, C.J.
West s.n., 7 Jul 1993, WELT L04404, Bell’s Flat, C.J.
West s.n., n.d. WELT L04402.
Comments Stevens (1987) records Ramalina luciae
from Africa, Sri Lanka, Indonesia, Australia and across
the Pacic to Fiji, Cook Islands and Tahiti. In the
Kermadec Islands, R. luciae has only been collected
from trees on Raoul Island – it seems to be absent
from the same types of habitats on the nearby Meyer
Islands and Herald Islets, where it is to be replaced by
R. pacica. Morphologically, R. luciae is very similar to
R. pacica, with both species having long dichotomous
branches and large round to ellipsoid soralia. Chemistry
easily separates the species, with R. luciae having the
sekikaic acid aggregate (K+ pink) and R. pacica having
salazinic acid in the medulla (K+ red). Ramalina luciae
tends to have longer branches between dichotomies
(branching points). Stevens (1987) notes that apothecia
in Australia are uncommon; they are described as lateral,
with the disc 2 mm diam. and concave. The spores of R.
luciae are ellipsoid to fusiform, straight or rarely curved
(10–)12–16(–18) × 4–5(–6) µm.
Ramalina meridionalis Blanchon et Bannister
Description (Fig. 9.) Thallus corticolous or saxicolous
(in New Zealand rarely corticolous), grey-green to
yellow-green, suberect to pendulous, 3–12(–200) mm
long, sparse to densely branched; branching mainly
dichotomous, often with narrow lateral branches, usually
more densely branched at the base where the holdfast
can produce several branches; branch width 1.0–2.0 mm,
basal branches rigid angular-terete to slightly attened,
apical branches terete-subterete, twisted and somewhat
nodulose, attenuate with ne apices which often break,
leaving a blunt tip; surface matt, pseudocyphellae always
present, numerous, linear, long, often giving a striate or
striate-nervose appearance; holdfast delimited; soredia
absent. Outer cortex indistinct or absent, chondroid
strands present in medulla. Pycnidia present. Apothecia
not seen in material from the Kermadec Islands.
Chemistry: Usnic acid (major), norstictic acid (major)
and ± connorstictic acid (minor). Medulla K+ red, C–.
Description based on Blanchon and Bannister (2002).
Northern Kermadec Islands: Herald Islets: One
specimen seen: Nugent Island, P.J. de Lange K1162, 18
May 2012, AK 330752. Southern Kermadec Islands:
Macauley Island: One specimen seen: Mt Haszard,
north-western seaward cliffs, P.J. de Lange K1159, 21
May 2012, AK 330766.
Comments: This species has been recorded from
New Zealand, Lord Howe Island and Norfolk Island
(Bannister & Blanchon 2003) so its presence on the
Kermadec Islands was not unexpected. It is evidently
scarce on the Kermadec Islands, though it is possible
that it has been overlooked through confusion with
the supercially similar R. australiensis. Ramalina
meridionalis is most reliably distinguished from R.
australiensis by the presence of norstictic acid (K+ red)
and the lack of extensive secondary branching towards
the apices.
Figure 9. Ramalina meridionalis, Macauley Island, Mt
Haszard (seaward clis) (P.J. de Lange K1159, AK 330766).
Scale Bar 20 mm. Photo: Dan Blanchon
Lichen notes from the Kermadec Islands. II. Ramalina

178
Ramalina microspora Kremp.
Description (Fig. 10) Thallus saxicolous, orange-brown,
caespitose, 10.0–17.5 mm high. Branching dense,
entangled, with many small side branches, anastomoses
visible between adjacent branches; branches solid,
narrow, terete, subterete to attened (particularly at
base), minor branch tips curled and slightly attenuate;
branch width 0.1–2.0 mm, surface pitted and uneven;
no pseudocyphellae seen; holdfasts delimited, multiple,
some blackened. Apothecia common, subterminal,
marginal, laminal and some geniculate, disc 0.2–2.5
mm in diam., concave to plane when very young with
thick thalline margin, plane to convex and occluded
when mature, obscuring margin. Ascospores 8 per
ascus, elliptical, 1-septate, 10–14 × 3–4 μm. Chemistry:
Divaricatic acid. Medulla K–.
Description based on Blanchon et al. (2012).
Northern Kermadec Islands: Raoul Island: One
specimen seen: W.R.B. Oliver s.n., 1908, n.d., WELT
L04263. Herald Islets: Two specimens seen: Dayrell
Island, P.J. de Lange K552, 18 May 2011, AK 330753;
Dayrell Island, P.J. de Lange K1282, 18 May 2011,
UNITEC 6953.. Southern Kermadec Islands:
Macauley Island: Five specimens seen: Southern
edge of plateau, W.R. Sykes 1029/K, 22 Nov 1970, CHR
162670; Crater Walls (Mt Haszard), P.J. de Lange K554,
21 May 2011, AK 330755; Mt Haszard, north-western
seaward cliffs, P.J. de Lange K555, 21 May 2011, AK
330756; western seaward cliffs, P J. de Lange K559, 21
May 2011, AK 330760; Mt Haszard, western seaward
cliffs, P.J. de Lange K571, 21 May 2011, AK 330767.
Curtis Island: Four specimens seen: West side of
island, W.R. Sykes 1038/K, 28 Nov 1970 CHR 162672;
Around fumaroles near summit, G.A. Taylor s.n., 10 Nov
1989, AK 257905; AK 257906; Chasm, Foodtown Bay,
G.A. Taylor s.n., 10 Nov 1989, AK 257914. Cheeseman
Island: One specimen seen: Summit rock outcrops,
southern side of island, P.J. de Lange K560, 24 May
2011, AK 325084. L’Esperance Rock: One specimen
seen: P.J. de Lange K1158, 26 May 2011, AK 330763.
Comments Blanchon et al. (2012) conrmed Ramalina
microspora as present on Curtis, Cheeseman and Macauley
Islands; the main islands of the southern Kermadec
Islands. Subsequent examination of specimens for this
paper, extends the Kermadec range south to L’Esperance
Rock, and also conrms the species as present in the
northern Kermadec Islands (cf. Sykes 1977), where it
is known from Raoul Island and from Dayrell Island in
the Herald Islets chain. This is supposedly a Hawaiian
Islands endemic whose range was recently extended to
Rarotonga in the Cook Islands (Blanchon & de Lange
2011). Its presence on Rarotonga and the Kermadec
Islands follows a pattern already noted by Renner and de
Lange (2011) for Lejeunea hawaikiana M.A.M. Renner
et de Lange and L. schusteri Grolle, minute, corticolous
liverworts, originally described as New Zealand
endemics and subsequently recorded from Raoul Island
and Rarotonga. A note of an epiphyllous gathering of it
reported by Sykes (1977: 48, CHR 211875) is actually R.
geniculata, and Ramalina microspora appears to be an
exclusively saxicolous lichen (see Blanchon et al. 2012),
characterized by its densely branched caespitose thallus
which is studded with subterminal and marginal apothecia
and a secondary chemistry of divaricatic acid. Two other
species recorded from the Kermadec Islands, Ramalina
geniculata and R. exiguella, could be mistaken for R.
microspora. However, Ramalina geniculata is inated
and perforate, whereas R. exiguella lacks perforations
and medullary secondary chemistry.
Ramalina pacica Asah.
Description (Fig. 11A, B) Thallus corticolous rarely
saxicolous, grey-green to yellow-green, erect or subpen-
dulous to pendulous, 20–110 mm long; branching
dichotomous, sparse to dense and intricate; branch width
0.5–2.5 mm, branches at, some marginal and apical
splitting; broad basally, narrowing towards apices; apices
attenuate and forked; surface matt or shiny, smooth,
often with linear pseudocyphellae; holdfast delimited;
soralia marginal and laminal, round to ellipsoid, margins
prominent. Apothecia not seen. Chemistry: Usnic acid,
salazinic acid (major), +/– protocetraric acid (minor/
trace). Medulla K+ red.
Description based on Blanchon et al. (1996)
Northern Kermadec Islands: Raoul Island: Five
specimens seen: W.R.B. Oliver s.n., 1908, AK 202677;
W.R.B. Oliver s.n., 1908, WELT L04265; Lava Point, A.E.
Wright 12491, 26 Nov 1994, AK 224394; North Terraces,
B.W. Hayward KE34, 25 Nov 1994, AK 224400; near Boat
Cove, P.J. de Lange K546, 16 May 2011, AK 330746.
Meyer Islands: Three specimens seen: North Meyer,
W.R. Sykes 675/K, 6 Dec 1966, CHR 161137; North
Meyer, P.J. de Lange K542, 12 May 2011, AK 330742;
Peter J. de Lange & Dan J. Blanchon
Figure 10. Ramalina microspora, Curtis Island, near
summit (G.A. Taylor s.n., AK 257906). Scale Bar 20 mm.
Photo: Dan Blanchon

179
North Meyer, P.J. de Lange K1157, 12 May 2011, AK
330768; South Meyer, P.J. de Lange K544, 13 May 2011,
AK 330744. Herald Islets: Two specimens seen: North
Chanter, North Summit, P.J. de Lange K548, 16 May
2011 AK 330748; Dayrell Island, P.J. de Lange K1283, 18
May 2011, UNITEC 6957. Southern Kermadec Islands:
Macauley Island: One specimen seen: Mt Haszard,
western seaward cliffs, P.J. de Lange K558, 21 May 2011,
AK 330759. L’Esperance Rock: One specimen seen:
Summit, P.J. de Lange K561, 26 May 2011, AK 330762.
Comments Ramalina pacica is a mainly subtropical-
warm temperate species, found from South Africa through
Southeast Asia, Australia, New Zealand and the Cook
Islands (Stevens 1987, Blanchon et al. 1996). On the
continental islands of New Zealand it is now in decline,
resulting in its current listing as Threatened by the New
Zealand Lichen Threat Listing Panel (de Lange et al.
2012). However, on the Kermadec Islands it is one of the
more common Ramalina species, though it is now mostly
seen there in the northern Kermadec Island group on the
less densely vegetated Meyer Islands and Herald Islets.
It now seems less common on Raoul Island (cf. Oliver
1912, Sykes 1977), possibly because the forest cover
there has increased in density following the eradication of
goats from that island in 1983, thus providing conditions
that do not favour this photophilous species (de Lange
& Galloway 2015). In the southern Kermadec Islands,
Ramalina pacica was collected from Macauley Island
and L’Esperance Rock, and these seem to be the rst
records of the species from the southern Kermadec Islands.
Some saxicolous specimens, e.g. L’Esperance Rock (de
Lange K561, AK 330762), have stunted and deformed
morphology and only traces of salazinic acid in TLC. On
the islands, Ramalina pacica can be mistaken for broad
forms of R. peruviana but that species has denser irregular
branching and the sekikaic acid aggregate, whereas R.
pacica has dichotomous branching and salazinic acid.
The species can be sympatric with R. luciae but they are
easily distinguished by their chemistry. Apothecia are
unknown in New Zealand and Stevens (1987) reports
them as uncommon in Australian material. They are
described by Stevens (1987) as marginal, disc 1–3 mm
diam., concave, plane or convex, edges sometimes
incised; spores fusiform or ellipsoid, straight or slightly
curved, 12–16(–20) × 4–6 µm.
Ramalina peruviana Ach.
Description (Fig. 12) Thallus corticolous, rarely
saxicolous, grey-green to yellow green, tufted, erect to
subpendulous, 20–60 mm long; branching subdichotomous
to irregular, intricate; branch width 0.15–1.3 mm,
branches at to subterete or terete, branches can be slightly
twisted, apices sharp; surface matt or shiny, smooth or
rugose, pseudocyphellae sometimes present; holdfast
delimited or diffuse in dense colonies; soralia punctiform,
numerous, laminal, marginal or apical, small brils often
produced from soralia. Apothecia rare, marginal, lateral;
disc 0.4–2.7 mm diam., concave to plane, often incised;
ascospores narrow, fusiform, straight or rarely curved,
(9.5–)14.5–17.0(–23.0) × 3.0–6.0 µm. Chemistry: Usnic
acid, homosekikaic acid (major), sekikaic acid (major),
Figure 11. Ramalina pacica. A. Raoul Island, Lava Point, narrow form (A. E. Wright 12491, AK 224394).
B. Herald Islets, North Chanter, broad form (P. J. de Lange K548, AK 330748). Scale Bar 20 mm. Photo: Dan Blanchon
Lichen notes from the Kermadec Islands. II. Ramalina

180 Peter J. de Lange & Dan J. Blanchon
ramalinolic acid (minor/trace), 4'-O-demethylsekikaic
acid (minor/trace), 4'-O-methylnorsekikaic acid (minor/
trace), +/- 4'-O-methylnorhomosekikaic acid (trace).
Medulla K+ pink.
Description based on Blanchon et al. (1996)
Northern Kermadec Islands: Raoul Island: Five
specimens seen: W.R.B. Oliver s.n., 1908, AK 33480;
Between Meteorological Station, Boat Cove and Fishing
Rock, R.C. Cooper 44128, Jun 1956, AK 304651; Above
farm, W.R. Sykes 679/K, 3 Jan 1967, CHR385511; Old
orchard behind Met Station, B.W. Hayward KE31, 25
Nov 1994, AK 224386; Sunshine Valley, P.J. de Lange
K545, 16 May 2012, AK 330745. Meyer Islands: Two
specimens seen: North Meyer, P.J. de Lange K541,
12 May 2011, AK 330741; South Meyer, P.J. de Lange
K543, 13 May 2011, AK 330743. Herald Islets: Three
specimens seen: North Chanter, North Summit, P.J. de
Lange K549, 16 May 2011, AK 330749; Napier, P.J. de
Lange K550, 17 May 2011, AK 330750; Nugent, P.J. de
Lange K551, 18 May 2011, AK 330751.
Comments: Ramalina peruviana is a widely distributed
species. Stevens (1987) lists it from Africa, Australia,
New Zealand, South America and several Pacic islands.
In New Zealand, it is widespread in the North and northern
South Islands, also the Chatham and Three Kings Islands
(Bannister et al. 2004), and in the North Island; next to
R. celastri probably the most commonly encountered
species in the genus. On the Kermadec Islands, it
seems to be restricted to the northern Kermadec Islands
where it grows on rocks and bark. On the Kermadec
Islands Ramalina peruviana could be mistaken for R.
australiensis or R. pacica but R. australiensis lacks
soralia and has no medullary chemistry, and R. pacica
has simple dichotomous branching and salazinic acid in
the medulla rather than the sekikaic acid aggregate.
Incertae sedis
We were unable to identify one W.R.B. Oliver specimen
from Raoul Island (WELT L04264) to species level due
to its fragmentary nature and poor state of preservation.
Additionally, we were unable to locate two other
specimens held at AK, AK 227550 and AK 310551, both
from Raoul Island.
DISCUSSION
This paper accepts eleven Ramalina species for the
Kermadec Islands, an increase of four (R. canariensis,
R. leiodea, R. meridionalis and R. microspora) on those
accepted earlier for the islands by Bannister and Blanchon
(2003), and Galloway (2007). Of the species recorded
here, only R. leiodea is new to the mycobiota of the
New Zealand Botanical Region, R. microspora having
been nally conrmed as present in the New Zealand
Botanical Region (sensu Allan 1961) by Blanchon et
al. (2012). The discovery of Ramalina meridionalis on
the Kermadec Islands represents an anticipated easterly
range extension of this species, which had hitherto
been recorded from the northern North Island of New
Zealand, Lord Howe and Norfolk Islands (Blanchon &
Bannister 2002). The number of species recorded for the
Kermadec Group (eleven) compares favourably with
that of mainland New Zealand (15, Galloway 2007) or
nearby Norfolk Island (eight, <http://www.anbg.gov.au/
abrs/lichenlist/IS_NORFOLK. html/, viewed 14 May
2012>) and Lord Howe Island (ve, McCarthy 2012).
In contrast the seeming low diversity of Ramalina on
Lord Howe Island is unusual. This may reect the more
closely forested condition of that island, which would
limit the range of habitats for photophilous Ramalina,
or the possibility that the lichen mycobiota of that island
has yet to be thoroughly explored.
In a pattern that reects island size, relief, habitat
and vegetation recovery, Ramalina is best represented
only on the islands of the northern Kermadec Islands.
With the exception of R. microspora, none of the
Ramalina species recorded from the southern Kermadec
Islands (R. geniculata and R. pacica) are widespread,
though on Macauley Island, R. pacica is locally
common in a small area. Nevertheless, aside from
the Meyer Islands and Herald Islets, Ramalina is not
especially conspicuous on Raoul Island either, quite a
departure from the comments made by Sykes (1977)
of R. pacica: ‘The long grey fronds of this species
are common on Rhopalostylis trunks on the Terraces of
Raoul’ which was not the case in 2011. Indeed, aside
from a few selected microhabitats, such as boulder
beaches, cliff faces and on the exposed branches of
trees, in the high canopy, or along ridgelines, none of the
Ramalina recorded from the northern Kermadec Islands
were common. While some, such as Ramalina celastri,
are inexplicably scarce. We suspect that the changes in
abundance of some Ramalina species recorded by Oliver
(1912) and Sykes (1977) from Raoul Island, and the
scarcity of others, reects the ongoing recovery of that
island’s vegetation following the removal of goats (by
1983), and, by 2002, cats and rats (see de Lange & Havell
2009). The forest on Raoul Island has greatly changed
in its structure since the rst lichen collections were
made, with a general thickening up of the understorey.
Ramalina species are generally photophilous, and it
seems probable that the past abundance of species such
as R. pacica (Sykes 1977) can be attributed to the
then open forested conditions created over much of the
island by feral goats. Now, 29 years after the last goat
was eradicated, such habitats exist only where local
Figure 12. Ramalina peruviana, Herald Islets, Napier Island
(P. J. de Lange K550, AK 330750). Scale Bar 20 mm.
Photo: Dan Blanchon

181
conditions have kept the vegetation open (e.g. coastal
shrubland at Coral Bay, on the exposed ridgelines and
cliff faces of Hutchison, and Smiths Bluff, and at Boat
Cove and Denham Bay), within the caldera where
volcanic activity has kept the vegetation cover open, or
in places where human activity has kept the forest open,
such as along the Airstrip of the Terraces.
ACKNOWLEDGEMENTS
The authors thank Jeremy Rolfe for providing Figure 1,
and for his general assistance with preparing all of the
gures for this paper. We also thank Francis Duff for
her thoughtful comments on a draft of this manuscript.
The senior author is grateful to Matt Jolly (skipper)
and the crew of the RV Braveheart for assistance
with all landings made on the Kermadec Islands; Tom
Trnski (Team Leader of the Kermadec Biodiscovery
Expedition 2011) for his organization of the expedition,
and support of the botanical investigations carried out
on those islands during May 2011; and Warren Chinn
for company in the eld. We thank Ewen Cameron
(curator) and Dhahara Ranatunga [Auckland Museum
Herbarium (AK)], Leon Perrie (curator) and Anthony
Kusabs [Museum of New Zealand Te Papa Tongarewa
Herbarium (WELT)], Wanda Daley (PDD) and Sue Gibb
(CHR) for promptly answering our queries and granting
access to specimens held under their care.
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Peter J. de Lange, Science and Policy Group, Private Bag 68908, Department of Conservation, Newton, Auckland,
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Dan J. Blanchon, Department of Natural Sciences, Unitec Institute of Technology, Private Bag 92025, Auckland,
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Article submitted: March 2012; revised and accepted: August 2012.